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Publication of Memoirs ceased with Vol. 55.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
VOL. 96. No.1
A STUDY OF THE SNAKE, TACHYlofENIS PERUVIANA
WIEGMANN AND ITS ALLIES
By WARREN F. \VALKER, .JR.
WITH FIVE P L A T E ~
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
NOVEMBER, 1945
No. 1. - A Study of the Snake, Tachymenis peruviana
Wiegmann and Its Allies
CONTENTS
PAGE
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Historical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Geographi c Conditions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Composition of peruviana . ........ . .... ' . . . . . . . . . . . . . . . . . . . . . . . . . 7
Localities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Acknowledgments ......................... ' .' . . . . . . . . . . . . . . . . . . . 14
The peruviana Complex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Tachymenis peruviana ,\\Tiegmann . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Tachymenis tarmensis spec. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21
Tachymenis affinis Boulenger. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 22
The attenuata Complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24
Tachymenis attenuata aUenuata. spec. et subsp. nov. . . . . . . . . . . . . . .. 24
Tachynwnis attenuata boliviana subsp. nov. . . . . . . . . . . . . . . . . . . . . .. 26
The chilens;s Complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 28
Tachymenis chilensis chileusis (Schlegel). . . . . . . . . . . . . . . . . . . . . . . .. 28
Tachymenis chilensis assimilis (Jan) ........... . ....... . .. : . . . .. 32
Tachymenis chiiHlnsis melanura subsp. nov.. . . . . . . . . . . . . . . . . . . . .. 35
Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 38
Relationship of the Described Forms. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 38
Outlying Spccies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 43
Redefinition of Tachy-menis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 44
Notes on the Habits of the Various Forms. . . . . . . . . . . . . . . . . . . . . . .. 47
Key to TachymeniB pcruviana and Allies. . . . . . . . . . . . . . . . . . . . . . . . .. 52
Relationship of Tachymenis to other Boiginae . . . . . . . . . . . . . . . . . . . .. 53
Bibliography 54
Plates . . . . ............................ ............... ...... .... 56
4 BULLETIN: MUSEUM OF COMPARAtIVE ZOOLOGY
INTRODUCTION
Historical
TachYlllenis was established by Wiegmann in 1835 on the basis of a
specimen from Peru. The type was named peruviana. Although
Wiegmann's characterization was adequate, many species from all
parts of the world were subsequently referred to the genus, which
naturally caused it to have very little real meaning. By the time of
Boulenger's Catalogue (1893-1896) Boettger, Boulenger, and Cope had
simplified the situation by placing the Old World forms in either
Tarbophis or Amplorhinus, and most of the New World ones in Ery-
throlamprus, as follows:
Tachymenis dromiciformis Peters 1863
Tachym.ni. vivax GUnther 1864
Tachymenis fissidens Peters 1869
Tachym.nis melanocephala Peters 1869
Tachymenis taeniata Peters 1869
Tachymenis piceivittis GUnther 1872
Tachy,nenis nototaenia Peters 1882
Tachymeni. bipunctata Garman 1883
Tachymeni. flssidens Garman 1883
Tachymenis imperialis Garman 1883
Tachymeni. lataitia Garman 1895
Tachymeni. dicipiens Gunther 1895
= Erythrolamprus dromicifo,..mis
(Peters) 1863.
= Tarbophi. savignyi (Savigny) 1829.
= Erythrolamprus imperialis (Baird
and Girard) 1859.
= Erythrolampru. lateritiu8 (Cope)
186!.
-= Erythrolamprus piceivittis (Cope)
1869.
= Erythrolamprus piceivittis (Cope)
1869.
=Amplorhinus nototaenia (Giinther)
1864.
=Erythrolamprus bipunctata (Gun-
ther) 1858.
= Erythrolamprus fi.sidens (Gunther)
1858.
= Erythrolamprus imperialis (Baird
and Girard) 1859.
= Erythrolamprus lateritius (Cope)
1863.
= Erythrolamprus dicipiens (Gun-
ther) 1895.
Tachymenis grammophrys Gunther 1895 =Erythrolampru. grammophrys
(Duges) 1888.
Boulenger completed the job in his Cata/{Jgue, and redefined l'achy-
men;s to include only peruviana, which he believed to range from cen-
tral Peru through Bolivia to southern Chile, and affinis of central Peru.
The latter was a new species which he described. He included Coron,lIa
WALKER: THE SNAKE, TACHYMENIS PERUVIANA WIEGMANN 5
chilensis Schlegel 1837 and Psammophylax assimilis Jan 1863 in
the synonymy of peruviana. All of this was a significant contribution
toward an understanding of the genus.
However, in following years large numbers of peruviana (as rede-
fined by Boulenger) were accumulated by the museums of the world.
These were taken at elevations ranging from sea level to the high
Andean plateaus of 13,000 feet, and from localities extending from
central Peru to southern Chile - a range in latitude equivalent to the
distance from New York to Panama! Naturally considerable geo-
graphic variation was encountered, and similarly a great deal of varia-
tion found in the specimens. Investigators began to feel that several
races or even species were involved. Werner (1898, 1901, 1904) recog-
nized five varieties of peruviana: p. peruviana in Peru,' p. vittata in cen-
tral Chile, p. coronel/ina in north central Chile, p. catenata in north
An indication of the climatic variation can easily be obtained from an examination of rainfal
and temperature figures. Data from Carlson (1943) for a few selected stations follow:
Station
Cuzco. Peru . ........... .
Arequipa. Peru . ........ .
Antofagasta, Chile . ..... .
Coquimbo. Chile .. ...... .
Santiago, Chile . ........ .
Valdivia, Chile ......... .
Puerto Montt, Chile . .... .
Annual
Rain/all
(Inches)
31.7
4.2
0.2
4.5
13.8
105.5
86.1
Period 0/
Heaviest
Rai1.
January
February
j ~ ~ ~ '
June
June
July
Annual
Average
Temp. (F)
51.3
56.8
63.0
58.3
57.0
52.9
51.8
Temperatur e
Ranoe
6.9
2.3
12,4
10.4
23.0
16,4
13.6
N.B. No specimens have been taken at Antofagasta. The record is merely included to indi-
cate the aridity of northern Chile. Specimens have been taken at all other stations.
central and central Chile, and p. dorsalis in Bolivia. These varieties
were poorly defined and, since they only made matters confusing, they
have not been generally recognized. Amaral (1930) followed Boulenger
in recognizing only peruviana and aJfinis in the area from central Peru
through Bolivia to southern Chile. Hellmich (1938), working primarily
with Chilean material, made another attempt to break up the apparent
vast array of forms included in peruviana, but came to the conclusion
that only one widespread and very polymorphic species was involved.
The primary purpose of this work has been to review the situation
and make a more careful study of the variations of peruviana (as rede-
fined by Boulenger) throughout its entire range, in order to deter-
mine whether or not more than one species is involved, or, if only one,
whether or not there is any clinal sequence in the variation. In clarify-
ing the status of peruviana, the generic type, I have also considered its
6
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
relation to ajfinis of Peru and to the three outlying species which have
been added to the genus since Boulenger's Catalogue: brasiliensis
Gomes (1918) from Sao Paulo, Brazil; elongata Despax (1910) from
northern Peru; and surinamensis Dunn (1922) from Surinam (Dutch
Guiana). Thus, in a sense, this paper constitutes a generic revision,
but, since no specimens of elongata and b-rasiliensis are available and
the descriptions are inadequate, my conclusions regarding the outlying
forms are tentative, and this study centers around peruviana. and its
immediate allies.
Geographic Conditions
In order to understand the morphological variation found in peru-
viana (sensu Boulenger) some knowledge of the geographic conditions
encountered by this snake in its wide range is necessary. The bulk of
the Peruvian and Bolivian specimens have come from the Andean
region from central Peru into Bolivia. Apparently they are primarily
highland snakes for they occur at least as high as 13,000 feet, and few
have been taken at low elevations. The lowest record on the west side
of the Andes is 1,630 meters (about 5,350 feet); on the east side a few
have worked their way farther down from the mountains and have
reached at least the edge of the tropical lowlands. Specimens have
been taken in the predominantly lowland department of ;\'Iadre de Dios
in Peru and also from the city of Santa Cruz (elevation about 500
meters, 1,640 feet) in Bolivia. Consequently many of these snakes
have encountered alpine conditions with a vegetation characterized by
procumbent dwarf shrubs and low cushion or rosette-shaped plants;
others, the more favorable and moist conditions found on descending
the eastern side of the mountains, where areas of grass steppes and
scattered shrubs, savanna and evergreen woods, and tropical rain
forest are encountered more or less in succession. Those from the
western slopes of the Andes in this region have found little precipita-
tion due to prevailing south-easterly winds. Thus the climate at the
higher elevations is semiarid with a xerophytic vegetation of cacti,
low shrubs, and heath. Lower down, there is a belt of desert crossed by
fertile river valleys. This is complicated along the narrow coastal
plain by a rich lomas vegetation dependant upon heavy fogs, but
Tachymcllis does not extend this low in Peru.
The Chilean specimens have been taken at lower elevations along
the coast and western slopes of the Andes in the central part of the
country from the province of Atacama to Chiloe. They are apparently
absent from the extreme deserts of northern Chile where nitrate de-
posits, salt basins, and intermittent rivers are the characteristic fea-
tures. Neither are they found in the cold damp forests of the far south.
Thus these snakes have avoided the extremes, but even in central
Chile they have encountered a wide range of topographic and climatic
conditions. The Andes tend to become progressively lower on going
south. At first the mountains ascend more or less directly from a nar-
row coastal plain, but to the south, a low coastal range of 1,000 feet
to 1,500 feet elevation and a broad central valley are interposed
between the Pacific Ocean and the Andes proper. In going south in
this region there is also an accompanying climatic change fully as
pronounced as that in topography. The temperature gradually de-
creases, and the prevailing winds, which are off shore in the north,
shift to come in from the ocean. This causes a pronounced precipita-
tion gradient. The northern deserts give way to semiarid conditions
in Atacama, and these to a mild mediterranean climate from the
southern part of the province of Coquimbo to Bio Bio. South of the
Rio Bio Bio forests appear and the climate is marine, a condition which
becomes very pronounced in the far south.
Composition of peruviana
Given the diversification of geographic conditions outlined above,
it is not surprising that peruviana, as hitherto considered, shows a
great deal of variation in coloration, squamation, dentition, and even
in penial characters. With the combined material of the American
Museum of Natural History, the Chicago Museum of Natural History,
the Museum of Comparative Zoology, and the United States National
Museum, enough specimens from all parts of the range were brought
together to be able to analyze the variation quite accurately. It was
soon obvious that peruviana was merely an omnium gatherum.
With such a large amount of material (209 specimens from 59 locali-
ties). it has been relatively easy to separate the forms involved. Un-
fortunately, however, some crucial regions where intergration might be
expected are poorly represented. Consequently, in breaking up the
array of forms previously included in peruviana, the question of which
are races and which species has been largely a matter of personal
judgment based on the degree of difference. I have no illusions as to
the permanency of my findings. Further revision will doubtless be
necessary as more material becomes available.
8 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
As I see the situation, peruviana can be broken down into three
species complexes. One found in the Andean regions of central and
southern Peru and Bolivia, a second along the edge of the tropical
lowlands of Peru and Bolivia, and a third along the coast of Chile.
These I have called the peruviana, attenuata, and chilensis complexes
respectively. Kames were available in the synonymy of peruviana for
many of the forms in these groups.
The peruviana complex includes pcruviana of Wiegmann and tar-
mensis to be described. I have also placed affinis of Boulenger in this
group. These species are characterized by a fairly stocky body form.
The body of the hemipenis is covered with spines of sub equal length,
its base only has a few minute ones, and its tip is covered by small
spines which fuse at their bases to give a slightly calyculate appear-
ance. There are 5-12 solid maxillary teeth. Normally they have 1 pre-
and 2 postoculars, and 8 supralabials with the fourth and fifth entering
the orbit.
The attenuata complex contains a new species with two races:
attenuata attenuata and attenuata bo/iviana. The body form of these is
quite elongate in comparison with members of the other complexes.
This is reflected in a greater number of caudals (60- 69 as opposed to
37- 55 in the other two groups), total ventrals (210-221 as opposed to
176-2(7), and ratio of tail length to total length (0.22-{).24 as opposed
to 0.14-{).21). The spines on the body of the hemiphenis increase in
size posteriorly, the base has several very large spines as well as
scattered minute ones, and the tip is distinctly calyculate with only
slight traces of spines. The solid maxillary teeth are more numerons
than in either other complex ranging from 12-16. The oculars are
variable, but the supralabials are still normally 8(4-5).
The chilcnsis complex contains three races arranged in north-south
sequence along the coast of Chile: chilensis assimilis (Jan), chilen,sis
chilensis (Schlegel), and chilensis melanura to be described. Jnall, the
body proportions, and consequently the ventral scales and ratio of tail
length to total. length, are close to those of peruviana, but the penis
resembles that of attenuata. The solid maxillary teeth range from 6-8.
There are typically 2 pre- and 2 postoculars, and the supralabials are
normally 7(3-4).
In summary peruviana as hitherto considered breaks up as follows:
The pcruviana complex:
perUl,;ana \Yiegmann
tarmensis spec. nov.
The attenuata complex:
attenuata attenuata spec. nov.
attenuata boliviana subsp. nov.
The chilensis complex:
chilensis chilensis (Schlegel)
chilensis assimilis (Jan)
chilensis melanura subsp. nov.
Boulenger's affinis appears to belong to the peruvialla complex.
These three complexes will be described in detail. This will be fol-
lowed by a discussion of their relationships to one another and to the
outlying forms of Tachymenis, my concept of the genus, and the habits
of the various forms.
Localities
In as much as South American collecting sites are frequently rather
unfamiliar towns, I shall give a brief description of the localities from
which specimens have been examined before proceeding. In this list, I
have followed the spelling on the American Geographical Society's
Map of Hispanic America where possible in an attempt to standardize
the spelling of localities.! Unless specific and accurate elevations were
available, those of the American Geographical Society's maps have
similarly been followed. The writings of Bingham (1916), Bowman
(1916), Stiglich (1922-23), and Weberbauer (1936) have been extremely
useful in obtaining information about Peruvian localities, and Carlson
(1943) and Rudolf (1929) for Bolivian and Chilean places. The loca-
tion of the department or province of any given locality can be deter-
mined by consulting the accompanying map of the distribution of the
various forms (PI. 1).
Peruvian Localities:
Arequipa: The capital city of the coastal department of Arequipa.
Its elevation is about 7,500 feet. The region is semiarid except in
irrigated valley bottoms.
Chospiyoc: A town in the lower part of the Cuzco valley south of
Ollantaytambo, department of Cuzco. Its elevation is 10,000
feet. The vegetation consists primarily of grassy steppes.
Chucuito: A town on the shore of Lake Titicaca just south of the
city of Puna, department of Puno. It is in a region of grass
steppes. The elevation is 3,820 meters.
1 In accordance with present American trends, and in the interest of simplicity, I have dropped
all the Spanish accents from the place names.
Collacachi: A town in the department of Puno, about five miles
southwest of Chucuito.
Cuajones: A mining camp located at an elevation of 3,230 meters
in the upper tributaries of the Rio Moquegua, department of
Moquegua. The region is semiarid.
Cuzco: The capital city of the department of Cuzco. The city is
located at an elevation of 3,495 meters in a broad fertile valley
surrounded by grassy highlands.
Huacullani: A small town in the southern part of the department
of Puno a bit northeast of Pisacoma. It is located at an elevation
of about 4,000 meters at the edge of the Titicaca basin very near
the Bolivian border. The vegetation is alpine.
Huanta: A large city in the northern part of the department of
Ayacucho. It is located at an elevation of 2,730 meters on a small
tributary of the lower Montaro in a region with a vegetation that
is transitional between the highland grass steppes and evergreen
woods.
H uarocondo: A town in the central part of the Cuzco valley between
Chospiyoc and Cuzco, department of Cuzco. Its elevation is
11,000 feet. The vegetation consists primarily of grassy steppes.
Huispang: A town in the department of Cuzco. It is probably in
the Urubamba valley northwest of the city of Cuzco. Its eleva-
tion is 12,175 feet.
Juliaca: A city in the department of Puno near the northwest end
of Lake Titicaca. Its elevation is 3,825 meters. The vegetation
consists primarily of grassy steppes.
Macchu Picchu: Famous Inca ruins located above the deep Toron-
toy gorge of the Rio Urubamba, department of Cuzco. Elevations
change rapidly in the region, but specimens from here are said to
have been taken at timberline at 12,000 feet elevation. The vege-
tation is transitional between that of the highlands and that of the
tropical lowlands.
Madre de Dios: A department of southeastern Peru. l\lost of it
belongs to the tropical lowlands but there are a few foothills of the
Andes in the southwestern part which reach a height of 2,500
meters. It is not known from what part of the department the
specimens came.
Ollantaytambo: A town in the Urubamba valley above the ruins
of Macchu Picchu, department of Cuzco. Its elevation is 9,400
feet. The vegetation consists of grassy steppes.
WALKER: THE SNAKE, TACIlYMENIS PERUVIANA WIEGMANN 11
Pachacayo: A town in the J\Iontaro valley between Oroya and
Jauja at an elevation of about 3,500 meters, department of Junin.
The vegetation consists of grassy steppes.
Pisacoma: A small town located at an elevation of about 4,000
meters in the southern part of the department of Puno. It is very
near Huacullani. The vegetation is alpine.
Pucara: Inca ruins in the department of Puno located near the
Cuzco railroad at an elevation of 3,880 meters. The vegetation is
alpine.
Puno: Capital city of the department of Puno located on the north-
western shore of Lake Titicaca at an elevation of 3,820 meters.
Puquiura: A town in the Cuzco valley located at an elevation of
9,500 feet between Cuzco and Huaracondo, department of Cuzco.
The vegeta tion consists of grassy steppes.
San Anton: A town in the department of Puno located in the north-
ern part of the Lake Titicaca basin at an elevation of 3,990 meters.
The vegetation is alpine.
Sicuani: A town in the department of Cuzco along the railroad to
the capital. The elevation is 3,515 meters; the vegetation alpine.
Tarma: A town in the department of Junin encountered on the way
to the Chanchamayo region. It is located near timber line at an
elevation of about 3,000 meters.
Toquepala: A mining camp located at an elevation of about 9,000
feet in the upper tributaries of the Rio Locumba, department of
Tacna. The climate is semiarid.
Urco: A hacienda in the upper part of the Urubamba valley above
the town of Urubamba, department of Cuzco. The elevation is
about 3,000 1lleters. The vegetation consists primarily of grassy
steppes.
Vitor: A town in the VitoI' valley below Arequipa, department of
Arequipa. Its elevation is 1,630 meters; the climate semiarid.
Yanama: A town in the Montara valley just above Jauja, depart-
ment of Junin. The elevation is about 3,500 meters. Tbe vege-
tation consists of grassy steppes.
Yauyos: A town in the upper tributaries of the Rio de Canete at an
elevation of 2,930 meters, department of Lima. The climate is
semiarid.
Yunguyo: A town in the department of Puno located along the
southern shore of Lake Titicaca at an elevation of 3,820 meters.
12 BULLETIN: MUSEUM OF ZOOLOGY
Yura: A town in the department of Arequipa. It is located a bit
to the northwest of the city of Arequipa at an elevation of 2,575
meters. The climate is semiarid.
Bolivian Localities:
Incachaca: A town somewhere in the department of Cochabamba.
Most of the department is below 3,000 meters, and has a vegeta-
tion transitional between that of the highlands and that of the
lowlands.
Santa Cruz de la Sierra: Capital city of the department of Santa
Cruz. The city is located at an elevation of about 500 meters
near the edge of the tropical lowlands.
Chilean Localities:
Angol: A city on the east side of the coastal range in the province
of Bio Bio. The mountains in the vicinity are under 1,000 meters.
The climate is slightly marine.
Cerros de Aculio: A local range of mountains somewhere in the
southern part of the province of Santiago. Their elevation is
probably under 2,000 meters. The climate is mediterranean.
Chanarcillo: A town about 40 miles from the coast in an intermit-
tent river valley of the semiarid province of Atacama. Its eleva-
tion is between 500 and 1000 meters.
Chile: A few specimens have no data other than the country. One,
with no data at all, is undoubtedly from Chile.
Chiquaihue: A town near Collipulli in the southern part of the
province of Bio Bio. Its exact location is not known, but Colli-
pulli is located at an elevation of 200 to 500 meters on the main
Andean range just west of the central valley. The climate is
slightly marine.
Concepcion: It has been assumed that specimens with this data
have come from the vicinity of the coastal city of Concepcion-
capital of the province of Concepcion. The climate is transitional
between mediterranean and marine.
Coracol: A small town on the coastal range in the province of
Concepcion. The mountains are under 1,000 meters in elevation;
the climate transitional between mediterranean and marine.
Curacautin: A town on the western slopes of the Andes in the
province of Cautin. Its elevation is about 500 meters; the climate
slightly marine.
Domeyko: A semiarid range of mountains in the northern part of
the province of Atacama. They reach a height of over 4,000
meters. It is not known from what part of the range the speci-
mens came.
El Tofo: A town located about 10 miles in from the ocean along the
semiarid coast of northern Coquimbo. Its elevation is between
500 and 1,000 meters.
El Vergel: A town located somewhere near Angol in the province
of Bio Bio. See Ango!.
Fundo Hualpencillo: A town somewhere near Concepcion in the
province of Concepcion. See Concepcion.
Las Condes: A town in the province of Santiago, but its exact
location is not known. See Santiago.
Las Tundras: A small town near Chillan, the capital of the province
of Nuble. Chillan is located in the central valley at an elevation
of less than 200 meters. The climate is mediterranean.
Mafil: A town located in the central valley of the province of Val-
divia at an elevation of less than 200 meters. It is along the rail-
road connecting Valdivia with Cautin. The climate is marine.
N"ahuelbuta Hills: Part of the coastal range west of Angol in the
province of Bio Bio. The mountains are under 1,000 meters
elevation. The climate is slightly marine.
Puerto Corral: An ocean port in the province of Valdivia just south
of the city of Valdivia. The climate is marine.
Puerto Montt: A port on the Gulf of Ancud in the northern part of
the province of Chiloe. The climate is marine.
Punta Teatino: A small village located along the semiarid coast of
Coquimbo just north of La Serena.
Santiago: The capital of Chile, located in the province of Santiago.
The mountains in the province reacb a height of about 5,000
meters. The climate is mediterranean. The city of Santiago is
located in the beginning of the central valley at an elevation of
about 500 meters. Some specimens with this locality are known
to have been taken near the city: others from elsewhere in the
department.
Talcaguano: A coastal village just north of the city of Concepcion
in the province of Concepcion. The climate is transitional from
mediterranean to marine.
Traiguen: A town in the central valley in the northern part of the
province of Cautin. Its elevation is under 200 meters. The cli-
mate is slightly marine.
Valparaiso: The chief sea port of Chile, being the port of entry for
Santiago. It is located in the southern part of the province of
Aconcagua. The climate is mediterranean.
Acknowledgments
At this time I wish to thank Mr. C. M. Bogert, Dr. D.l\L Cochran,
!\Ir. A. Loveridge, and Mr. C. H. Pope for the loan of the material
which made this study possible, and for allowing me a free hand in its
utilization. I also wish to express my deepest gratitude to Dr. T. Bar-
bour, !\Ir. A. Loveridge, and Mr. K. P. Schmidt for suggestions and
help during the course of these investigations, and to Dr. F. M. Car-
penter for assistance in making the photographs.
The PERUVIAKA Complex
TACHYMENIS PERUVIANA 'Viegmann
Plate 2, fig. 2; Plate 3, figs. 5-9.
1835 Tachymenis peruviana Wiegmann, N. Acta. Ac. Leop.-Carol., 17, p. 252,
pI. 20, fig. 1 (type 10cality=Peru). Ophis peruviana Fitzinger, 1843,
Syst. Rept., pt. 1, p. 28. Tachymenis peruviana Peters, 1863, Monatsb.
K. Preuss. Akad. Wissensch. Berlin, p. 275 (in part).' Boulenger,
1896, Cat. Sn. Brit. Mus. Nat. Hist., 3, p. liS (in part: Aschiquiri,
Bolivia; Larecaja, Bolivia; Lake Titicaca). Barbour and Noble, 1921,
Proc. U.S. Nat. Mus., 58, p. 618 (i;' part: Pucyura
2
at 9,500 feet,
Chospiyoc at 10,000 feet, Huarocondo at lI,OOO feet, Ollantaytambo
at 9,400 feet - all localities in the department of Cuzco, Peru).
Escomel, 1929, Fauna de Arequipa, p. 9. Amaral, 1930, Mem. Inst.
Butantan Sao Paulo, 4 (1929), p. 210 (in part). HeIlmich, 1938,
Rev. Chilena Hist. Nat., 41, p. 108 (in part: Bolivian localities-
Carquayeoleo near Rio Muleto in the department of Uyuni,' Cerro
Sapo in the department of Cochabamba, Ayopayo at 3,200 meters
in the department of Cochabamba). Schmidt and Walker, 1943,
Zool. Ser. Field Mus. Nat. Hist., 24, pp. 290, 315 (in part: city of
Arequipa in Pcru and some of the other specimens believed to be from
the department of Madre de Dios in Peru).'
1845 Ophis peruana Tschudi, Fauna Per., Herp., p. 58 (nomem nudum.
Tschudi had no specimens. He was merely verifying Fitzinger's
placement of peruviana into Ophis, but misspelled the name).
I After this had gone to preHS my attention was called to another reference to T. peTUiliana
as follows; Garman, 1875, Bull. . 3, p. 278 (Cuzco Valley).
2 This town is spelled "Puquiura" on the maps of the American Geographical Society.
3 See footnote I, p. 20.
, One other reference to Tar:hymenis peruviana in t he literature results from a misidentification
of the specimen: Amaral, 1925, Proc. U.S. Nat. 67, Art. 2-1, p. 14 (U.S.N.M. 12277 from
Guayaquil, Ecuador = Dryophylax nattereri Mikan).
1854 Dipsas chilensis Dumeril and Bibron (not of Schlegel) Erpet. Gen., 7,
p. 1159 (in part).
1901 l'achymenis peruviana var. dorsalis Werner, Abhandl. und Bcrich. Kon.
Zool. Anthrop.-Ethnog. Mus. Dresden, 9(2), p. 8 (type locality =
Bolivia).
1915 Leimadophis andicolus Barbour, Proc. BioI. Soc. Washington, 28, p.
149 (type 10cality=Huispang at 12,175 feet, department of Cuzco,
Peru).
Wiegmann (1835) does not give a specific type locality for peruv,:ano"
so it is here restricted to the city of Puna in the department of Puna.
The choice of Puna is based on the fact that Dr. F. J. F. Meyen, who
collected the type specimen during his trip around the world in 1830-32,
mentions visiting this city in an account of h i ~ travels (1834). More-
over, specimens from Puna agree very well with the figure and descrip-
tion of the type. The type was probably a female judging by scale
count and tail shape. Because of the importance of peruviana as the
type of the genus, it is redescribed in detail on the basis of specimens
from Puna. These are then compared with specimens from other parts
of thc range to determine the amount of variation.
lJI aterial Examined. A total of 111 specimens (55 Q, 56 ci') refer-
able to peruviana have been studicd. A list of the localities from which
they carne follows:
Peruvian Localities:
Department of Arequipa
Arequipa: 1 Q, I ci' (C.M.N.H. 34055, 340(1)
Vitor: 1 ci' (C.M.N.H. 342(0)
Yura: 2 ci' (C.l\I.N.H.34201-2)
Department of Ayachucho
Huanta: 6 Q, 6 ci' (C.l\I.N.H 39647-58)
Department of Cuzco
Chospiyoc: 1 ci' (U.S.N.l\I. (0740)1
Cuzco and the Cuzco Valley: 4 Q, 3 ci' (C.l\LN.H. 40216-19,
l\I.C.Z. 3584-86)'
Huarocondo: 4 Q, 2 ci' (M.C.Z. 12412-3; U.S.N.l\I. 60697-8,
(0722)1
Huispang: 2 ci' (M.C.Z. 10986-87)'
Ollantaytambo: I Q (U.S.N.l\I. 60743)1
Puquiura: 2 Q, 1 ci' (U.S.N.M: 60681-3)1
Sicuani: 1 Q (A.l\I.N.H.36154)
Urea: 4 Q, 3 ci' (C.l\J.N.H. 34105-07, 34131-33, 34142)
Department of Junin
Pachacayo: 2 ,, 2 r:J' (M.C.Z. 42435-38)
Yanama: 1 r:J' (l\r.C.Z.45916)
Department of Lima
Yauyos: 1 ,, 1 r:J' (M.C.Z.45914-15)
Department of Madre de Dios: 3 ,, 4 r:J' (C.M.N.H. 40070,
40073-78)1
Department of Moquegua
Cuajones: 3 ', 4 r:J' (l\f.C.Z.46651-5i)
Department of Puno
Chucuito: 1 ,, 1 r:J' (C.M.N.H.40214-15)
Collacachi: 2 ,, 4 r:J' (C.M.N.H.34156-61)
Huacullani: 1 ' (C.M.N.H.40213)
Juliaca : 1 ' (C.M.N.H.34086)
Pisacoma: 1 ' (C.l\f.N.H.40212)
Pucara: 1 ,, 2 r:J' (M.C.Z. 45903-{)5)
Puno: 2 ,, 3 r:J' (A.l\LN.H. 36155, C.M.N.H. 34259, U.S.N.lVf.
109015- 17)'
San Anton: 2 ', 1 r:J' (A.l\f.N.H. 5256, C.M.N.H. 4065-66)
Yunguyo: 11 ,, 10 r:J' (C.l\LN.H.40174- 94)
Department of Tacna
Toquepala: 1 r:J' (l\LC.Z.45941)
Bolivian Localities:
Department of Santa Cruz
Santa Cruz de la Sierra: 1 ' (A.l\f.N.H.8793)
Diagnosis. This species is the type of the genus Tachymenis, and
forms the basis of the pCTuviana complex. The characters of this com-
plex have been discussed in the introduction. T. peruviana differs from
other members of the group chiefly in having 19 midbody scale rows,
dorsal scales with single apical pits, and generally a distinctly spotted
color pattern.
Description of W]Jotypcs. Fi\'e topotypes ha\'e been examined. All
are from the city of Puna in the department of Puno. These specimens
are A.M.N.H. 36155, a male collected by Beck; C.l\I.N.H. 34259,
also a male collected by K. P. Schmidt on October 28,1939; U.S.Xl\L
109015- 17, one male (109016) and two femal es collected by J. Soukup.
All are snakes of moderate size. The body is rather stocky and cylin-
drical with a more or less oval shaped head that is only slightly distinct
from the neck. The eye is of moderate size with a pupil that is sub-
elliptical in shape. The hemipenis is bifurcate with a bifurcate ductus
spermaticus. Its tip is covered with small spines which fuse at their
bases giving a slightly calyculate appearance. The body of the hemi-
penis is covered with larger spines of subequal size. There are no greatly
enlarged basal spines, but there are a few scattered minute ones. The
tbree topotypes whose dentition was examined have 6 solid maxillary
teeth wbich are subequal or increase slightly in size posteriorly. These
are followed after a gap by a pair of enlarged grooved fangs. The man-
dibular teeth decrease in size posteriorly after the first few.
The rostral is broader than deep, slightly visible from above; inter-
nasals a bit shorter than the prefrontals; frontallY2 times as long as
broad, longer tban its distance from the end of the snout, as long as the
parietals; supraocular of normal size; nasal entire or semi-divided;
loreal as deep as long or a bit deeper; 1 preocular reaching tbe top of the
bead but not contacting the frontal; 2 postoculars in three of the speci-
mens, 1 in the other two; temporals 2-3 in half the cases, 1-2 or 1-3 in
the others; supralabials 8(4-5); infralabials 9 in most of tbe specimens
with the first four in contact with the anterior pair of chin shields
which arc as long or longer than the posterior pair, but combinations
of 8(3) and 10(5) were also found.
The dorsal scales are smooth, have single apical pits, and a reduction
pattern that is typically 19-19- 15, but one specimen has only 14 pre-
anal rows. The abdominals are rounded, 141- 147 in females and 144-
147 in males; anal divided; caudals paired, 43-45 in females and 47-51
in males; total ventrals 186-190 in females and 191-198 in males.
The dorsal ground color of tbese snakes is light brown or grey dusted
with black. In addition, most of the dorsal scales bear a single black
apical dot which is about the size of and in the general position of the
apical pit. There is a short light middorsal line on tbe neck. This
narrows posteriorly and breaks up into a series of small middorsal
spots. There is a short black stripe on either side of the midcervical
line. Posteriorly this too breaks up into a series of spots. These spots
are of moderate size and dark. Those of opposite sides alternate a bit
and some fuse across the midline forming faint zigzag markings.
Laterally, on scale rows four and five, there is another series of dark
spots which fuse posteriorly to form a dark lateral line. The head
1 Some of these have already been discussed in the literature which is listed in the
synonymy.
2 Type and paratype of Lei madophis andicolu8 Barbour 1915. This specimen was reidentified
as Tachyme1,is peruvianu by Barbour and Noble in 1921.
3 Topotypes of peruviana.
18 BULLETIN; MUSEUM OF COMPARATIVE ZOOLOGY
coloration is also brownish or grey with irregular dark markings. A
small black line runs from the snout through the eye to the angle of the
mouth. It is best developed behind the eye. A similar line extends
from the parietals to the angle of the ja w. There is a small black mark
under each eye. The supralabials may also have a few other dark mark-
ings, and the infralabials and gulars a few scattered ones as well. The
ventral ground color is light grey or yellow. The abdominals and
caudals are covered with irregular dark brown or black markings.
The markings in the center of the abdominals and caudals are scattered,
but laterally they tend to form a line.
The largest topotype, a female, has a total length of 375 mm.; tail
65 mm. The ratio of tail length to total length is 0.16-D.17 in the
female topotypes; and 0.18 in the males.
Variation. The remaining material agrees very well with the topo-
types as regards major characters, but differences in. detail are com-
mon. The body form, penial characters (fig. 2), and dentition are the
same in all. The number of solid maxillary teeth ranges from (i- 10
and averages 8 in twenty specimens which have been examined from
various parts of the range. The type was said to have 5 such teeth.
There are also some differences in the proportions and number of
cephalic plates. The internasals are either as long as or shorter than the
prefrontals; frontal 1 ~ to 2 times as long as broad, longer than its
distance from the end of the snout, about as long as the parietals but it
deviates a bit in either direction; nasal single or semidivided, rarely
divided. Generally there is only 1 preocular, but 2 are present in J 6
head halves. Typically there are 2 postoculars, but these fuse with
one another or with a supralabial to form 1 in 18 head halves. The
temporal formula is 2-3 in the majority of cases, but variations are
common. Combinations of 1-2, 1-3,2-2, and 2-4 occur in 65 head
halves. Nearly all of these combinations could easily have been de-
rived from a basic 2-3 pattern. The supralabials typically have a
formula of 8(4-5). However, two of the first labials have fused in 13
head hakes to give a combination of 7(3- 4), one other specimen has a
count of 8(3-4-5) on one side of the head, and in two additional cases
a preorbitallabial has divided to give a formula of 9(5-6). There are
usually 9 infralabials, the first 4 or 5 of which contact the anterior
pair of chin shields, but variations of 8(3), 10(4), 10(5), and 1O(i1)
occur in 28 head halves.
The dorsal scales in all specimens are smooth and ha'e a single
apical pit, but the pit is very poorly developed in certain populations.
Only a small amount of variation is encountered in the reduction
pattern of the dorsal scales. One specimen has 17 cervical rows and
two others have 20. Only one deviates at midbody with a count of 18.
Four specimens have 16 or 17 preanal rows, but in half of these the
number is reduced to the typical 15 a bit posteriorly. Two specimens
have as few as 14 preanal rows. The number of female abdominals
ranges from 134-152 with an average of 144; males 136-158, average
J 48. The anal plate is divided in all, and the left portion generally
overlaps the right, but the reverse is the case in two mutant individuals,
and there is a partial fusion of the two halves in another specimen.
The female caudal range is 37-50, average 42; males 44-55, average 49.
These caudals are typically paired, but in eight cases 2-6 were single.
The female total ventral range is 176-196, average 186; males 184-209,
average 197. Scale counts reported in the literature lower the male
abdominal and caudal range to 135 and 37 respectively, but do not
otherwise affect the above observations.
The majority of the specimens are quite similar to the topotypes in
general color pattern, but may differ in a few details. The typical
coloration can be seen in figures 5- 7. Variations occur in the row of
light middorsal spots which fuse to form a wavy line in many speci-
mens. Other deviations are the failure of any of the large dorsal spots
to fuse across the midline; the reduction of the series of lateral spots;
more perfuse dark markings on the labials; and a nearly complete
absence of dark markings on the abdominals and caudals. Young
specimens have a coloration similar to that in typical adults.
Aside from the above departures, several distinct color phases are
present in the material at hand. About one half of the }1adre de Dios
specimens are so melanistic that the typical pattern can only be de-
tected with difficulty (fig. 9) . The specimens from Puquiura are also
quite dark and in addition have a third dark diagonal line on the head.
This one extends from the first supralabial to the middle of the infra-
labials. In specimens from Yauyos (Lima), Chospiyoc (Cuzco),
Huispang (Cuzco), and Bolivia the light middorsal line and dark
lateral lines are very pronounced, and the dark dorsal spots are a bit
larger than usual (fig. 8). As near as can be told from the description,
this race was first noticed in.Bolivia by Werner (1901) and described
by him as dorsalis. However, its reappearance in scattered localities
shows that the race is not tenable. It would similarly be unwise to
name the other color varieties for they occur with or near typical
members, and are irlentical with them in other characters. The only
conclusion that can be reached is that peruviana is very polychromatic.
The females examined have an average total length of 368 mm.,
20
BULLETIN: MUSEUM Of' COMPARATIVE ZOOLOGY
tail 59 mm.; in males the total length averages 405 mm., tail 72 mm.
The largest female was taken at Urco and has a total length of 626 mm.,
tail 99 mm. The largest male is from Vitor and has a total length of
656 mm., tail III mm. The ratio of tail length to total length ranges
from 0.14--D.18 in the females, and from 0.16-0.21 in the males.
Runge. From the above data it is apparent that peruviana is a snake
of the Central Andes. Its range extends from the departments of
Junin and Lima in central Peru south along the Andean chain to south-
ern Bolivia.
l
Actual localities at which the species has been taken can
be found in the list of literature and material examined. It is a fairly
common snake, but appears to be most abundant in the Andes of
southern Peru. This, however, may only be a result of more extensive
collecting in that area. It has reached an elevation in southern Peru
of nearly 4,000 meters at several points: Huacullani, Pisacoma, San
Anton. It is thus essentially a highland form, but it does extend from
the upland valleys and plateau down on both sides of the Andes. The
lowest point it is known to have reached on the west coast is at Vitor
(1630 meters), but it has been taken as low as Santa Cruz in Bolivia
(c. 500 meters) on the east side. The vegetation in all of these regions
consists primarily of grass steppes as will be discussed later.'
Remarks. As can be seen from the above discussion, Tachymcnis
peruv;anu is a wide spread form even when it is considered in the re-
stricted sense used in this paper. Its range extends along the Andean
chain for nearly 1,000 miles. Most of the specimens in this area are
found in the highland valleys, but the species can apparently migrate
to a certain extent from one valley to another. This seems to. be
enough to scatter the variation and keep the species intact. Because
of the semi-isolation of each population, the variation is widely scat-
tered and not arranged in any elinal sequence. This is characteristic
of species in mountainous regions. Each population difl'ers from neigh-
boring ones noticeably in both squamation and coloration, and some,
such as those from Puquiura and Madre de Dios, differ enough to be
regarded as incipient races. However, it is dangerous to name such
forms. Ta.chymenis pcruvia.na would be an excellent form for the study
of speciation in mountainous areas. Although 11 J specimens have
I The southernmost record appears to be Carquaycoleo (Hellmich, 1938). I can not find thi8
town, but it is said to be near Uio :il.luleto in the department of Uyuni. No department, of Uyuni
is shown on the American Geographical Society's maps. The only ciue I can find to the location
of Carquayeolco isa town of Uyuni in the department of Potosi. This town is near a Rio ),Iuiatos.
2. After this was '\\.itten, my attention was caned to a specimen from Born Jesus de Lapa,
Bahia. Brazil (Carnegie ::\1useum 347) which was identified by Griffin (1916) as Tachymt' nis
peruviana. 1 have reexamined this specimen and agree with t he identification, but am skeptical
as to the accuracy of the locality record.
WALKER: THE SNAKE TACHYMENIS PERUVIANA WIEGMANN 21
been examined, most are from southern Peru. The other parts of the
range are poorly known.
TACHYMEl'lS TAR'iENSIS spec. nov.
Plate 4, figs. 10, 11.
Type. C.J\I.N.H. 5698, an adult female from Tarma in the depart-
ment of Junin, Peru. It was collected by M. P. Anderson on :May 19,
1914.
Diagnosis. It is a Tachymenis closely related to pcruviana, but
differing in having a few more solid maxillary teeth (12 as opposed
to a maximum of 10 in peruviana), a few more caudals (55 as opposed
to a maximum of 50 in females of peruviana) which causes the ratio of
tail length to total length to be higher, absence of scale pits, and a very
light dorsal ground color with practically no trace of a pattern.
Description of type. The body form, pupil shape, and dentition are
quite similar to the corresponding characters of peruviana. There are
12 solid maxillary teeth.
The rostral is broader than deep, visible from above; internasals
about as long as the prefrontals; frontal twice as long as broad, longer
than its distance from the end of the snout, as long as one and shorter
than the other parietal; supraocular about the size of the frontal;
- nasal entire; loreal a bit longer than deep; 1 preocular; 2 postoculars;
temporals 2-2; supralabials 8 (4- 5); infralabials 9(5) on the right side
and 8(4) on the left; anterior chin shields longer than the posterior.
The dorsal scales are smooth, lack apical pits, and have a reduction
pattern of 19-19-15. The abdominals are rounded, 135; anal divided;
caudals 55, 3 being single the rest paired; total ventrals 190.
The top of the head and body is nearly uniform greyish brown in
color (figs. 10-11). There is no trace of a pattern except for a bit of
black at the base of a few of the dorsal scales. The color of the under-
side is grey stippled with black. This stippling becomes heavier pos-
teriorly.
The snake has a total length of 393 mm.; length of tail 83 mm.;
ratio of tail length to total length 0.21.
Remarks and Range. This form resembles pcruviana in general
structure. It differs only a bit in such quantitative characters as num-
ber of teeth, number of caudals, and in the ratio of tail length to total
length. Its chief departure is in the absence of scale pits which are
often very poorly developed in pcruDiana, and in its unusual coloration.
Using this degree of difference and its geographic position at the
northern part of the range of peruviana as criteria, one would expect
tarmensis to be a race of peruviana. Perhaps it is, but until males be-
come available to cheek on penial characters it is best considered as
a full species belonging to the peruviana complex. It is probably a
rather obscure form, for it is only represented by the type from Tarma,
and the Tarma region is quite well known.
TACHYMENIS AFFINIS Boulenger
Plate 4, fig. 12.
1896 Tachymenis affinis Boulenger, Cat. Sn. Brit. Mus., 3, p. 319, pI. 7, fig. 1
(type locality = Muna', Peru). Amaral, 1930, Mem. lnst. Butantan
Sao Paulo, 4 (1929), p. 209.
1921 Tachymenis pernviana Barbour and Noble (not of Wiegmann), Proe.
U.S. Nat. Mus., 68, p. 618 (part: Cedrobamba ruins
2
at timber line,
12,000 feet, department of Cuzco, Peru).
Material. Tachymenis affilli. has previously been known by the
single type in the British Museum. One additional specimen turned
up during an examination of material from the National Museum.
This specimen, U.S.N.M. 60721, an adult male, was collected at the
Inca ruins of Macchu Picchu in the department of Cuzco, Peru by
E. Heller in June 1915 while he was a member of the Kational Geo-
graphic Society - Yale University Peruvian Expedition. It was first
identified as peruviana by Barbour and Noble (1921) in their report of
the Expedition's reptiles. But since it has 17 midbody scale rows and
a reduced color pattern, there is little doubt but that it is affinis. This
specimen enlarges our knowledge of this rare species. It is described
and compared with the type below.
Diagnosis. It is a Tachyrnenis differing from all other members of
the genus in having a dorsal scale reduction pattern of 17-17-15 in-
stead of 19-19-15. It further ditrers from peruviana in having a re-
dnced color pattern, bnt resembles it so much in other features that
it has been included in the peruviana complex.
Description. T. affinis is similar to peruviana in body form, penial
characters, and general dental pattern. The specimcn in question has
9 solid maxillary teeth that are subequal in size and are followed after
1 StigJich (922) cites two localities with this name in Peru - one along the northern coast of
Ancash, and one on the upper part of the Rio Huall aga in Hualluco. ] am not sure which is the
type locality but it is probably the latter since the latter is the larger of the two and is in the
highlands (2,000-2,500 meters).
discussed below. The Ccdrobamba ruins are now better
a gap by a pair of enlarged grooved fangs. The type had 15 solid
maxillary teetb. This may seem like a great discrepancy, but the
maxillary range in peruviana is at least 5- 10.
The cephalic plates of the specimen examined are quite similar to
those of peruviana except that the frontal is only 173' times as long as
broad, and is shorter than the parietals. The nasal is semidivided.
There is 1 preocular; 2 postoculars; temporals 2-2 on the right side,
shrivelled on the left; supralabials 8(4-5); infralabials 9(5). The
cephalic plates in the type were essentially the same. Its temporals
were 2-3.
The dorsal scales are smooth, have single apical pits, and a reduc-
tion pattern of 17-17-15. The abdominals are rounded, 147; anal
divided; caudals 50, 6 of these are single and the rest paired; total
ventrals 197. The type, also a male, had 153 abdominals, 57 caudals,
and 2lO total ventrals.
As can be seen, the specimen of affinis from Macchu Picchu agrees
very well with the type in the above characters, but it does differ a bit
in coloration. Both are brownish above, hut the dorsal scales are
darker than the lateral ones in the Macchu Picchu specimen (fig. 12)
while the lateral scales are said to be darker in the type. However, I
do not believe that this is too significant for the color pattern of both
specimens seems to have been derived from a condition similar to that
in pcruviana by a process of reduction of the prominent dark stripes
and spots. Both specimens have fair remnants of the head markings
of peruviana. The type has traces of the rows of dark dorsal spots.
These are obscure in the Macchu Picchu specimen, but traces of the
light middorsal line can be seen. Both lack the black apical dots that
are so prominent in the dorsal scales of peruviana. The ventral scales
of the Macchu Picchu specimen are darker than in the type.
The snake examined has a total length of 454 mm.; tail 84 mm.;
ratio of tail length to total length 0.18. The type was a bit larger, and
had a ratio of 0.21.
Remarks and Range. T. affinis appears to be a very rare form of the
highland valleys of Peru whose range is known to extend from Muna
in Huanuco to Macchu Picchu in Cuzco. It must consequently over-
lap a large part of the range of peruviana al though the two have not
yet been taken together. This fact together with the marked differ-
ence in the reduction pattern of the dorsal scales precludes the possi-
bility of the two forms being races of the same species. There is no
doubt that affinis is a separate species, but I have placed it in the
pcruviana complex since the two are so close in penial characters.
The ATTEN"UATA Complex
TACHYMENIS ATTENUATA ATTENUATA spec. et suhsp. nov.
Plate 2, fig. 3; plate 4, figs. 13-15.
1943 Tachymenis lJefuviana Schmidt and Walker (not of Wiegmann), Zoo!.
Ser. Field Mus. Nat. Hist., 24, p. 290 (in part: some of Peruvian
specimens).
Type. C.M.N.H. 40071, an adult male collected by Sr. Juan Perea,
and originally deposited in the University of Arequipa. The specimen
has no further data than Peru, but such material in the University's
collection is believed to be from the department of Madre de Dios
(Schmidt and Walker, 19-13) ..
Diagnosis. It is a l'achymenis differing from members of the peru-
viana complex primarily in its more attenuate body proportions. This
results in a greater number of caudals and total ventrals, and in a
greater ratio of tail length to total length. Other differences can be
found in the condition of the hemipenis, and in the dentition. All of
these have been discussed in the introduction. This race differs from
bolitiana, to be descrihed, in having a slightly lower number of solid
maxillary teeth (12-14 as opposed to 14-16), and of abdominals and
caudals (in males 148-150 and 60-64 as opposed to 152 and 69), and
in having a speckled instead of a checkered color pattern.
Description of type. The body form is quite attenuate in comparison
with peruviana; subeliptical in section. The head is moderately dis-
tinct from the neck. The eye is of average size with a subcliptical
shaped pupil. The hemipenis (fig. 3) is bifurcate with a bifurcate
ductus spermaticus; tip distinctly calyculate with only slight indica-
tions of spines at the top of the folds; body spinose with the length of
the spines increasing posteriorly; base with a row of 4 very large spines
followed by a few scattered minute ones. There are 12 solid maxillary
teeth which increase slightly in size posteriorly and are followed after
a gap by a pair of enlarged fangs; mandibular teeth decrease in size
posteriorly after the first few.
The rostral is broader than deep, visible from above; internasals
shorter than the prefrontals; frontallYz times as long as broad, longer
than its distance from the end of the snout, distinctly shorter than the
parietals; supraocular of normal size; nasal entire; loreal deeper than
long; 1 preocular extending to the top of the head but not contacting
the frontal; 2 postoculars; temporals 2- 2; supralabials 8(4-5); 8 in-
WALKER: THE SNAKE, TACnYMENIS PERUV[ANA WIEGMANN 25
fralabials with the first 5 in contact with the anterior pair of chin
shields which are as long as or longer than the posterior pair.
pattern of 19-19-15; abdominals rounded, 150; anal divided; caudals
paired, 60; total ventrals 210.
The ground coloration above and below is a light brown which is so
heavily stippled and flaked with black that the general appearance is
dark grey. However, upon careful examination traces of a pattern
similar to that of per1lviana can be seen beneath all of this. (See figs.
13-15). These traces are especially prominent on the head and neck,
for a diagonal black line runs from the eye to the corner of the mouth,
a second one extends from the parietals to the angle of the jaw, and
there is a distinct black stripe on either side of the midline on the
back of the neck. A small black line also runs along the sides of the
belly but becomes obscure posteriorly for the general black stippling
becomes very dense.
The specimen has a total length of 582 mm.; tail 12G mm.; ratio of
tail length to total length 0.22.
Notes on paratlJpe. On paratype, C.l\LN.H. 40072, also an adult
male has the same data as the type. It is also quite similar to the type
in general body form, penial characters, and dentition. It has 7 large
basal spines' on the hemipenis, and 14 solid maxillary teeth. The
cephalic plates are the same with the exception of minor differences:
2 preoculars; temporals 2-3 on the right side and 2-2 on the left; in-
fralabials 8(5) on the right side and 9(5) on the left. The dorsal scales
have the same reduction pattern, but have traces of two apical pits!
There are 148 abdominals, G4 caudals, and 214 total ventrals. The
greatest departure is in coloration. The head, neck and belly are simi-
lar to the type, but the back is a nearly uniform light greyish brown
with a minimum of black stippling and flaking. Its general appearance
is close to that of tarmensis. The paratype has a total length of 581
mm.; tail 131 mm.; ratio of tail length to total length 0.23.
Remarks and Range. The slight indication of scale pits in the para-
type is puzzling. This fact together with the aberrant color might
warrant the recognition of this specimen as a separate form, but this
does not seem advisable at this time for the two resemble each other in
all other characters, and Tachymenis is known to be very polychromatic
and to show some variation in terms of scale pits.
It is unfortunate that more specific locality data are not available
for this race. Both this form and perl/viana are found in the depart-
ment of Madre de Dios. Its more elongate body form suggests the
possibility that it is a tropical lowland type, while peruviana may come
from the mountains in the southern part of the department.
Morphologically, a. attenuata resembles tarmensis in the greater
number of solid maxillary teeth, lack of scale pits (at least in the type),
and greater number of caudals. But this does not necessarily indicate
relationship. T. tarmensis is presumably a local northern form so the
two are separated by a large geographic hiatus. Furthermore, al-
though the coloration of a. attenuata is variable it is nearer to that of
peruviana than to that of tarmensis. Its resemblances to -tarmensis
could easily have been arrived at through parallel evolution. It seems
more probable that both attenuata and tarmensis evolved indepen-
dently from peruviana stock. -
TACHYMENIS ATTENUATA BOLIVIANA subsp. nov.
Plate 4, fig. 16.
Type. A.J\I.N.H. 36020, an adult female taken at an elevation of
2,500 meters at Incachaca in the department of Cochabamba, Bolivia
by J. Steinbach.
Diagnosis. It is a Tachymenis whicb resembles a. attenuata very
closely. ' As mentioned above, it differs primarily in coloration, but
tbis is supplemented by slight differences in the number of solid maxil-
lary teeth, abdominals, and caudals.
Dese-ription of type. The general body form, pupil shape, and denti-
tion are the same as in a. attenuata. There are, however, 16 solid
maxillary teeth.
The proportions of the cephalic plates are also the same. There are
2 preoculars; 2 postoculars; temporals 2-3; supralabials 8(3-4-5); in-
fralabials 10(5) on the right and 9(4) on the left.
pattern of 19- 1\i-15; abdominals rounded, 149; anal divided; caudals
paired, 61; total ventrals 210.
The color pattern (fig. 16) is closer to the color of peruviana than is
that of a. attenuata. The dorsal ground color is light brown. On either
side of the light middorsal line, there is a row of fairly large uniformly
shaped dark spots. Those of opposite sides alternate with one another,
fuse with one another at their corners across the midline, and also
alternate with and fuse with a less distinct row of lateral spots. The
net result is a checkered pattern which is especially prominent an-
teriorly. The various rows of spots can be easily homologized with
WALKER: THE SNAKE, TACllYMENIS PERUVIANA WIEGMANN 27
similar rows in peruviana, but the spots are not as large and generally
do not fuse in peruviana. The underside is a light brown flaked with
black. Anteriorly these black markings are arranged into four longi-
tudinal lines, but posteriorly they merge with the darkened ground
color. The head and neck markings are the same as in typical peru-
viana.
The type has a total length of 576 mm.; tail 125 mm.; ratio of tail
length to total length 0.22.
Notes on paratypes. Two paratypes A.:vr.N.R 36015 (female) and
A.lVI.N.H. 36017 (male) have the same data as the type. The female
specimen resembles the type very closely. It has 14 solid maxillary'
teeth. There is 1 preocular; 2 postoculars on the left side and 1 on the
right; temporals 2-3; supralabials 9(5-6); infralabials 9(4) on the left
side and 9(5) on the right. There are 156 abdominals; 65 caudals, and
221 total ventrals. The coloration is the same as in the type. The
specimen has a total length of 630 mm.; tail 142 mm.; ratio of tail
length to total length 0.23.
The male specimen is also quite close. It has 16 solid maxillary
teeth. Its hemipenis is similar to that of a. attenuata, and has 7 large
basal spines. There is 1 preocular; 2 postoculars; temporals 2-3;
supralabials 8(4-5); infralabials 9(5). The dorsal scales, however, are
unusual in that there is an indication of a single apical pit. There are
152 abdominals; 69 caudals; 221 total ventrals. This specimen is very
melanistic, so only traces of the checkered pattern and head-neck mark-
ings can be seen. Its total length Is 661 mm.; tail 162 mm.; ratio 0.24.
Remarlcs and Range. As in a. attenuata the indication of scale pits
and the aberrant coloration of one of the paratypes presents a problem.
It is probably merely an example of the variability of these characters,
but more material is needed to be sure. The presence of a pit in a
minority of the population may indicate the recent derivation of this
form from pitted ancestors.
There is little doubt in my mind that typical attenuata and boliviana
are two races of the same species despite the present lack of inter-
grades for they have so many features in common. As a matter of
fact, their separation is merely a function of coloration (a variable
character) supplemented by minor differences in teeth and scale
counts. Both races are probably found along the edge of the tropical
lowlands of the Amazon basin - attenuata being a northern race and
boliviana a southern one.
28 BUI_LETIN: MUSEUM OF COMPARATIvE ZOOLOGY
The CHILENSIS Complex
TACHYMENIS CHILENSIS CHILENSIS (Schlegel)
Plate 2, fig. 4; plate 5, figs. 17-19,21.
1837 CoroneUa chilensis Schlegel, Phys. Serp., 2, p. 70 (type locality = Chile).
Guichenot, 1854, in Gay, Hist. Chile, Zool. 2, p. 79, pI. 4, fig. 1
(Valdivia and Colchagua, Chile). Dipsas Ghilensis, Dumeril and
Bibron, 1854, Erpet. Gen., 7, p. 1159 (in part: Chile). Tadlymenis
chilensis Girard, 1854, Proc. Acad. Nat. Sci. Philadelphia, p. 226
(Santiago, Chile). Girard, 1855, U. S. Naval Astron. Exped., 2, p.
213, pI. 37, figs. 1-6 (Santiago, Chile). Gunther, 1858, Cat. Colubr.
Sn., p. 34 (Colchagua and Tichitata in Chile, and Chile). Cope, 1860,
Proc. Acad. Nat. Sci. Philadelphia, p. 247 (Talcahuano
1
and Quin-
quina ld., Chile). Mesotes chilensis Jan, 1863, Arch. Zool. Anat.
Fisiol., 2, p. 308 (Chile). Jan, 1866, Icon. Gen. des Ophid., 18, pl. 6,
fig. 2 (Chile).
1863 Tachymenis perumana Peters (not of Wiegmann), Monatsb. K Preuss.
Akad. WissenRch. Berlin, p. 275 (in part). Steindachner, 1867, Novara
Rept., p. 62 (Chile). Boulenger, 1896, Cat. Sn. Brit. Mus. Nat. Hist.,
3, p. 118 (in part: Colchagua and Talcahuana
l
in Chile, southern
Chile, Chile). Werner, 1896, Verhandl. der KK Zool.-Bot. Gesellsch.
Wien, 46, p. 356 (Frutillar, Chile). Amaral, 1930, Mem. lnst. Butan-
tan Sao Paulo, 4 (1929), p. 210 (in part). Hellmich, 1938, Rev.
Chilena Hist. Nat., 41, p. 108 (in part: Puerto Montt in Chile, and
some other Chilean specimens). SchIDidt and Walker, 1943, Zool. Ser.
Field Mus. Nat. Hist., 24, p. 315 (in part).
1898 Tachymenis peruviana var. mllata' Werner, Zool. Jabrb. Sup., 4, p. 259,
pl. 13, fig. 9c (type locality = Frutillar, Chile). Werner, 1904, Ergeb.
Hamburger Magalh. Sammelr., in N aturhistorishes Mus. zu Hamburg,
1, Rept. und Batr., p. 15 (Coronel, Valdivia, and Puerto Montt, all in
Chile).
1898 Tachymenis affin;s Werner of Boulenger), Zool. Jahrb. Sup., 4, p.
259, pl. 13, fig. 9 d (Chile).
1904 Tachymenis perumana var.catenata' Werner,Ergeb. Hamburger Magalh.
Sammelr., in Naturhistorishes Mus. zu Hamburg, 1, Rept. und Batr.,
p. 14 (Estancilla in Valdivia, Chile).
Although Schlegel (1837) based his very brief description of chilensis
on two specimens collected by M. Lesson and 1\1. Garnot from an un-
known part of Chile, the scale count of these snakes and their essen-
I This locality is spelled "TalcagQsno" on maps ,of the American, Geographical Society.
2 These specimens were first ,reported by \Ve,rner tn l 896 as .. See above. .
3 This variety was first described by 'Verner 1111898 on the baSIS of specimens from CoqUlmbo.
The type series. judging from locality and coloration, should be referred to c. assimilis. The
present series, however. appears to he c. chilensis.
WALKER: THE SNAKE, TACHYUENIS PERUVIA:-IA WIEGMANN 29
tially lineate color pattern warrant the restriction of this name as a
trinominal to the most common race of the chilc1l81:s complex. This
race occurs in central and part of southern Chile. In as much as it has
been poorly defined and is very common, it is redescribed in detail on
the basis of the material examined.
Matcrial examined. A total of 57 specimens (26 <;1,31 ci") referable
to this race have been examined. All are from Chile. A list of the
localities from which they came follows. The provinces are arranged
in a north-south sequence.
Chile- no further data: 3 <;1,3 ci" (C.M.N.H. 23815; M.C.Z. 1978,
2539,3692; U.S.N.M. 436, 56442)
Province of Aconcagua
20 miles northeast of Valparaiso: 1 <;I (C.:\1.N.H. 40117)
Province of Santiago: 1 <;1,2 ci" (C.l\I.N.H. 9943, 9945-46)
In or near Santiago: 3 <;I, 1 ci" (M.C.Z. 2058; U.S.N.I\J. 5514,
3 specs.)
Cerro de Aculeo: 1 <;I. 1 ci" (I\I.C.Z. 19976- 77)
Province of Nuble
Las Tundras: 1 <;I, 1 ci" (l\I.C.Z. 6513, 2 specs.)
Province of Concepcion
Concepcion: 2 <;1,5 ci" (l\I.C.Z. 16372-74, C.l\I.N.H. 3863-66)
Coracol: 1 <;I, 1 9 (A.M.N.H. 18328, 18333)
Fundo Hualpencillo: 1 <;I (C.:\I.N.H. 31616)
Talcaguano: 5 <;1,7 ci" (A.I\I.KH. 21159; l\1.C.Z. 2065 - 2 specs.,
2066 - 2 specs., 2540 - 4 specs., 2766)
Province of Bio Bio
Angol: 1 ci" (C.M.N.H.23812)
Chiquaihue: 2 <;I (C.l\l.N.H.23808-{)9)
EI Vergel: 1 ci" (C.l\1.N.H. 31615)
Nahuelbuta Hills: 3 <;1,2 ci" (C.M.KH. 23810- 11, 23813-14, 31614)
Province of Cautin
Curacautin: 4 ci" (C.J\1..:-l.H. 6221-24)
Traiguen: 1 9 (C.J\f.N.H. 31617)
Province of Valdivia
Puerto Corral: 1 <;I, 1 ci" (A.M.KH. 36149- 50)
Province of Chiloe
Puerto ~ I o n t t : 1 ci" (A.I\1.N.H.27659)
Diagnosis. A Tachymrnis which resembles l'eruviana in general body
proportions and consequently in scale counts, but resembles attenuata
in penial characters. Its supralabials are normally 7(3-4). This form
has been placed into a separate species complex for these reasons as
30 BULLETIN: OF COMPARATIVE ZOOLOGY
has been pointed out in the introduction. This particular race differs
from the other two in the chilcnsis complex primarily in having a dorsal
color pattern that is distinctly Iineate,yet the dark dorsallinesarequite
narrow (not over 1 scale wide). T. c. ehilellsis also tends to have more
caudals, and total ventrals than either assimilis or
mcia1!'Ura..
Descriptio11. These snakes are of the samc ordcr of size as perul,iana,
have similar body proportions, and pupil shape. The hemipenis
(fig. 4), however, resembles that of atfenuata. It is bifurcate with a
bifurcate ductus spermaticus; tip distinctly calyculate with remnants
of minute spines on the tips of the folds; body spinose with the spines
increasing in length posteriorly; base with a row of very large spines
followed by a few scattered minute ones. The number of large basal
spines ranges from 4-5 in the specimens studied. The solid maxillary
teeth are subeqllal or increase slightly in size posteriorly. They are
followed after a gap by a pair of enlarged grooved fangs. The number
of solid maxillary teeth ranges only from 6- 7 in the material at hand,
bllt one specimen figured in the literature (.Jan, 1866) had 10 such
teeth. The mandibular teeth decrease in size posteriorly after the
first few.
1
The rostral is broader than deep, slightly visible from above; inter-
nasals shorter than the prefrontals; frontal lY;l- I% times as long as
broad, a bit longer than its distance from the end of the snout, shorter
than the parietals in the adults, but as' long as the parietals in the
young; supraoeular of normal size; nasal entire, rarely semi-divided;
IQreal' longer than deep or as long as deep; preoculars 2, rarely 3, the
upper one reaching the top of the head but not contacting the frontal;
postoeulars 2, rarely 1 or 3; temporals usually 2-3 but variations of 1-2,
1-3, and 2- 2 are very common, and combinations of 1-1, 2-4, and 3-2
occur; supralabials normally 7(3-4), but 9 head halves have 8(4-5)
and one has 8(3-4); infralabials generally 9 with the first four or five
contacting the anterior pair of chin shields which are as long as or
longer than the posterior; but combinations of 8(3), 8(4), 10(4), and
10(5) are also found, Aside from one specimen with a single preoeular,
reports in the literature do not increase the above variation,
The dorsal scales are smooth, have single well developed apical pits,
and a reduction pattern that is typically 19-19-15. A bit of variation
is encountered in the number of preanal rows, Ten specimens have 16
o'r 17 preanal rows which are reduced to 15 posteriorly, two specimens
have 17 rows which are not reduced, and one specimen has only 14
preanal rows. The abdominals are rounded, female range 148-158,
average 152; male range 152- 1G9, average 159. The anal is divided
The caudals are typically paired, but in two cases several are single,
female range 37-51, average 43; male range 45-53, average 49. The
total ventral range in females is 185-204, average 194; males 197-217,
average 208'. Scale counts reported in the literature lower the female
caudal rangc to 33 and the male caudal range to 39, but do not other-
wise affect the above obsenations.
Even though the coloration of c. chilcnsis is extremely variable,
certain common features can he detected, and similarities with the
pattern of peruviana be seen. In the majority of cases (fig. 17 and 21)
the ground color is a light brownish-grey, and most of the dorsal scales
bear a black apical dot about the size of and in the position of the apical
pit. The light middorsal line is vcry pronounced, but there are only
. traces of the prominent rows of black spots which bordered it in peru-
viana. The spots have been reduced and merely consist of a series of
black streaks which are never more than one scale wide. These fuse
into a narrow line. In some specimens they are completely lost in the
posterior regions. As in pl'r1l-viana, there is another black line on the
fourth row of scales, but in c. chilensis, this is accentuated by a slight
lightening of the ground color immediately above it, and a general
darkening of the ground color below it. The head has the typical
Tachymenis pattern consisting of a dark dash under the eye, a hlack
line extending from the eye to the angle of the mouth, and a similar
line extendi)lg from the parietals to the angle of the jaw. However,
the latter line is somewhat reduced as compared with peruvinnrt. The
color on the under side is yellowish with numerous irregular hlack
marks. Sometimes these are arranged in a linear manner. J\Iarkings
are nearly absent on the undersidc of the chin and tail.
The coloration of very young specimens is essentially the same as
just described. But the rows of black dorsal spots or streaks are a bit
more pronounced, and it is more apparent that the dark lateral line is
composed of a series of fused spots. Young from as far south as Puerto
Montt are spotted, but this is not as obvious in their case as in speci-
mens from central Chile.
Tn a number of specimens (fig. 18), the ground color above and
below is quite dark, and the light middorsal line is sharply set off hy
very distinct black lines. These lines are still only one scale wide.
This is the most striking departure, but it is connected to members with
the typical pattern by intermediate forms. In a few specimens, the
opposite has occurred (fig. 19). The ground color has becOlne very light
above the dark lateral line, and all traces of the dark dorsal lines ha\'e
been lost. Consequently the light middorsal line tends to blend with
the ground color and is not too distinct.
The females have an average total length of 318 mm., tail 50 mm.
The largest female, from an unknown part of Chile, has a length of
550 mm., tail 90 mm. Males averages 378 mm. overall, tail 65 mm.
The largest male, taken at Concepcion, has a total length of 597 mm.,
tail 95 mm. In the females, the ratio of tail length to total length is
0.14-0.17; in males 0.15-0.18. A couple of aberrant males, however,
extend the range of the latter to 0.13-o.33!
Range. T. c. chilensis is one of the more common snakes of central
Chile. Its range extends from the southern part of the province of
Aconcagua as far south as Puerto Montt 'in the province of Chiloe.
This includes the mediterranean region of central Chile and some of the
forested regions of the sOHth. The race seems to be more abundant in .
the former area. From the little data available on the vertical distribu-
tion of c. chilensis, it appears that it extends from sealevel to at least
2,000 meters elevation. It reaches the highest points in the northern
part of its range.
Remarks. Geographically, this race occurs between assimilis to the
north and melanura in parts of the south. It is also intermediate in
characters. The cline in the number of abdominals, caudals, and total
ventrals is lowest in assimiiis, reaches a maximum in chilensis, and falls
off a bit in mclanura as will be seen. The counts of ass inti lis are the
closest to peruviana. Similar tendencies ' can be seen in the coloration
change; northern assimilis is closest to peruvian a, chilensis has a
pattern which merely carries the differences in assimilis a step further,
and melanura is the farthest removed from peruviana. Evidence of this
sort strongly indicates that the entire chilclIBis complex has been de-
ri ved from
T ACHYMENIH CHILENSIS ASSIMILIS (.J an)
Plate 5, fig. 20.
1858 Tachymenis chilensis Girard (not of Wiegmann), U. S. Explor. Exped.,
20 Herp., p. 173 (Valparaiso, Chile).
1863 Psamnwphylax assimilis Jan, Arch. Zool. Anat. Fisiol., 2, p. 311(type
locality unknown). Jan, 1866, Icon. Gen. des Ophid., 18, pI. 1, fig. 2
(Chile).
1896 Tachymenis peruviana Boulenger (not of Wiegmann), Cat. Sn. Brit.
Mus. Nat. Hist., 3, p. 118 (in part: Coquirnbo, Chile). Amaral, 1930,
Mem. lnst. Butantan Sao Paulo, 4 (1929), p. 210 (in part). Hell-
mich, 1938, Rev. Chilena Hist. Nat., 41, p. 108 (in part: some of
Chilean specimens). Schmidt and Walker, 1943, Zoo!. Ser. Field
Mus. Nat. Hist., 24, p. 315 (in part).
1898 Taehymenis peruviana var. eoroncllina Werner, Zoo!. Jahrb. Sup., 4,
p. 259, pI. 13, fig. 9b (type locality = Coquimbo, Chile). Werner, 1904,
Ergeb. Hamburger Magalh. Sammelr., in Naturhistorisches Mus.
zu Hamburg, 1, Rept. und Batr., p. 14 (Valparaiso, Chile).
1898 Taehymenis pcruviana var. ealcnala Werner, Zoo!. Jahrb. Sup., 4, p.
259, p!. 13, fig. 9 a (type locality = Coquimbo, Chile).
In the original citation of assimilis (1863) Jan did not know the type
locality, and gave a very brief description of the animal. Fortunately
however, he subsequently (1866) figured this form on the .basis of a
specimen from Chile.! It is quite clear from this figure that the name
assimilis should be restricted to the most northern race of chilcnsis.
This race is redescribed below on the basis of specimens examined.
1\! aterial Examined. Sixteen specimens (7 "" 9 ei") have been
studied. A list of the localities from which they came follows:
No data: 1 ei" (AJI'1.N.H.37942)2
Chile: 1 ei" (C.i\tN.H. 7004)
Province of Atacama
Chanarcillo: 1 '" (l\f.C.Z. (514)
Domeyko: 1 '" (C.1I.N.H.5771)
Province of Coquimbo
El Tofo: 3 ",,2 ei" 36084-88)
Punta Teatino: 3 ei" 64939-41)
Province of Aconcaqua
Valparaiso: 2"" lei" (A.I1I.N.H. 36146-47, U.S.N.M. 5531)
Province of Santiago
Las Condes: 1 ei" (M.C.Z. 21211)
Diagnosis. It is a Tachymenis clearly belonging to the chilensis
complex, but differing from the typical race in having a slightly lower
number of abdominals, caudals, and total ventrals; and in having a
color pattern that is partly spotted and partly lineate.
Description. This race resembles chilensis very closely as regards
general proportions, hemipenial characters, and dentition. The number
of large, basal penial spines ranges from 2-6. There are six or seven
solid maxillary teeth followed after a gap by a pair of large, grooved
fangs.
The proportions and the disposition of the cephalic plates is also the
1 The figured specimen may be the actual type, for it agrees with the type dC8cription as
regards certain abtj.ormalities in the head shields.
2 This specimen closely resembles those from Coquimbo.
34 BULLETIN: MUSEI1M OF COMPARATIVE ZOOLOGY
same as in chilensis except that the frontal is a bit nearer the length of
the parietal. There are two pre oculars in every case except for one side
of a specimen from Punta Teatino in which the lower preocular has
fused with the loreal; postoculars 2; temporals generally 2- 3, but a
variation of 2-2 is common; supralabials 7(3-4); infralabials generally
9 with the first 4 or 5 in contact with the anterior pair of chin shields
which are as long as or longer than the posterior pair, but combinations
of 8(3) were found.
The dorsahcales are smooth, have single well developed apical pits,
and a reduction pattern of 19-19-15. The abdominals are rounded,
female range 141-148, average 146; male range 137-154, average 145.
Anal divided. Caudals paired, female range 38-44, average 41; male
range 40-49, average 46. The total ventral range in females is 181-190,
average 187; males 184-210, average 194.
Records in the literature extend the above variation a bit but not
significantly. The figure by .Jan (1866) shows 9 solid maxillary teeth
and supralabials 8(4-5). The type had both a single preocular and
postocular. Counts of Boulenger (1896) extend the female
caudal range to 49, and counts of Werner (1898) would extend their
abdominal range to 149.
The color pattern of c. assimilis (fig. 20) is intermediate between the
spotted pattern of p. peruvialla and the lineate one of c. chilclIsis. As a
rule, the dorsal ground color is a light brownish-grey, and most of the
scales have a black apical dot. The light middorsal line is present,
but it is frequently broken up into a series of middorsal spots. Anter-
iorly this line is bordered on each side by a row of fairly large dark
spots. Each row begins as a dark streak on the back of the neck. This
is the pattern of p. pcruv;alla; however, these spots fuse into a line at
about the middle of the body. In some cases they may even be fused
into a line for the entire length of the body thus ghing the lineate
appearance of c. chilclIsis. But if this is the case, the resulting line is
always more than one scale wide. There is also a distinct black lateral
line on the fourth row of scales as is for TachYl1lell is. The color
of the head and underside is the same as in c. chilellsis. The most com-
mon departure from the above was the blackening of the edges of many
of the dorsal scales. This partially obscured the typical pattern. In a
couple of other cases, the dorsal ground color was "ery light and the
pattern reduced.
The females examined have an average total length of 31G mm., tail
49 mm. The largest specimen, taken in Valparaiso, measures 510 mm.
overall, tail 80 mm. The males have an average total length of 372
WALKER: THE SNAKE, TACHYMENIS PERUVHNA 35
mm., tail 66 mm. The largest specimen was from Las Condes. It has
a total length of 484 mm. and a tail length of 81 mm. The ratio of tail
length to total length ranges from 0.15-0.16 in females; O.l6-{).19 in
males.
Remarks and Range. The range of assimilis extends from the
Province of Atacama in northern Chile south into the PrO\'ince of
Santiago. Thus it is primarily a north-central race, but it extends far
enough south to overlap slightly the range of chilcnsis. Both races are
found in the Provinces of Aconcagua and Santiago; however, assimilis
appears to be more common in Aconcagua and chilcnsis in Santiago.
To my knowledge, these two races have not been taken at the same
locality in this area of overlap. At the northern part of its range,
assimilis seems to be separated from the Peruvian and Bolivia forms
by the se\'ere deserts of the northern parts of Chile and southern
Bolivia, but more collecting in these regions is necessary to be certain.
The climate encountered by assimilis is primarily semiarid, but this
race does enter the mediterranean regions of Chile. This race has been
collected primarily along the coast, but does extend into the moun-
tains. It has been taken in thc Domeyko range, which reaches an
elevation of 4,000 meters, but we do not know from what part.
TACHYMF,NIS CHILENSIS MELANURA subsp. nov.
Plate 5, fig. 22.
Type. C.M.N.H. 3874, an adult male collected by C. C. Sanborn
between February 14 and 28, 1923 at Mafil, Province of Valdivia,
Chile.
Diagnosis. A Tachymenis belonging to the chi/ensis complex, but
differing from the typical race in ha\'ing such a great development of
melanin that the ground color is very dark and the typicallineate pat-
tern obscure. The race also tends to have a lower number of abdomin-
als, but this difference is not too significant.
Description of type. The general body shape is the same as in most
of the forms of Tachymcllis - of moderate size, cylindrical in section,
and an oval shaped head that is only slightly distinct from the neck.
The eye is of moderate size with a slightly subeliptical-shaped pupil.
The hemipenis is similar to typical chilensi.! being bifurcate with a
bifurcate ductus spermaticus; tip distinctly calyculate with small
spines on the tips of the folds; body covered with spines that increase
in size posteriorly; and 5 very large basal spines followed by a few
scattered minute ones. There are 8 solid maxillary teeth which increase
a bit in size posteriorly. These are followed after a gap by a pair of
enlarged grooved fangs. The mandibular teeth decrease slightly in
size posteriorly after the first few.
The rostral is broader than deep. visible from above; internasals
shorter than the prefrontals; frontal 172 times as long as broad. longer
than its distance from the end of the snout. distinctly shorter than the
parietals; supraocular of normal size; nasal entire; loreal a bit longer
than deep; 2 preoculars. the upper one reaching the top of the head but
not contacting the frontal; 3 postoculars; temporals 2-3 on the left
side. 2- 2 on the right; supralabials 7(3-4); 9 infralabials with the first
4 in contact with the anterior pair of chin shields which are longer than
the posterior pair.
The dorsal scales are smooth. have a well developed apical pit. and
a reduction pattern of 19- 19-15; abdominals rounded. 163; anal
divided; caudals paired. 52; total ventrals 215.
The dorsal ground color (fig. 22) is a uniform light black. This
naturally tends to obscure any pattern. but upon close examination
the remains of the typical chilensis one can be seen. There is a faint
indication of the light middorsal line bordered by a darker black. and
the narrow black line on the fourth row of scales. The latter still has a
slight lightening of the ground color just above it. The head too is very
dark. but the typical black streaks from the eyc to the angle of the
mouth and from the parietals to the angle of the jaw. as well as a few
dark supralabial markings can still be seen. The ventral color is a dark
grey with irregular black markings showing through. These tend to be
arranged in a linear manner. The undersides of the tip of the tail and
chin are much lighter. There are a few dark markings on the infra-
labials.
The type has a total length of 394 mm . tail 67 mm.; ratio of tail to
total length 0.17.
!I'otes on parai.'lprs. Seventeen paratypes (8 "'.9 <3') are available
(C.M.N. H. 3867-73. 3875-83. 5769). All are from Mafil and bear the
same data as the type. This series agrees very well with the type.
The number of large basal spines on the hemipenis ranges from 5- 8;
the number of solid maxillary teeth from 7-8. Most have 7.
The frontal is 172 to 1% times as long as broad. generally shorter
than the parietals but in the younger specimens the two plates are
subequal in length; nasal entire. rarely semidivided; loreal either longer
than decp or deeper than long; generally 2 preoculars. occasionally 3;
generally 2postoculars. occasionally 3; temporals usually 2-3 or 2-2.
but variations of 1-1. 1-2. 1-3. and 3-2 occur; supralahials 7(3-4) in
WALKER: THE SNAKE, TACHYMENIS PERUVlANA WIEGMANN 37
most cases, but one specimen has 6(3-4) and another 8(4-5); infrac
labials 9 ~ 4 ) Or 9(5) in most, but combinations of 8 ~ 3 ) , 10(4), and 10(5)
are present in this series; anterior chin shields generally longer than
the posterior, but they are the same length in It few cases.
The dorsal scales have a well developed apical pit and a typical re-
duction pattern of 19-19-15. Two specimens have 16 and 17 preanal
rows. In these cases the number is not reduced below 16. Abdominals
147-155 in females, average 150; males 146-163, average 158. Anal
divided. Caudals typically paired, but several specimens have 1-3
single; female range 41-51, average 45; males 40-54, average 49. The
total ventral range in females is 192-201, average 196; males 195-215,
average 207.
Ninety-five per cent of the paratypes have a coloration that is
practically identical with that of the type. These are all melanistic,
but vary a bit in the degree with which the underlying chilensis pat-
tern shows through. The melanism is absent in only one specimen
(C.M.N.H. 5769). This one consequently has the typical chilensis
pattern, but is referred to melanura on the basis of locality. No very
young specimens are present in the series, but judging from a late
embryo removed from C.;\,LN.H. 3878, the ground coloration would
be lighter and the lineate pattern more evident.
The females have an average total length of 444 mm., tail 72 mm.;
the largest specimen measures 497 mm. and 90 mm. The males have
an average total length of 422 mm., tail 72 mm.; the largest measures
568 mm. and 77 mm. The ratio of tail length of total length ranges
from 0.15-{).18 in the females; 0.15-{).18 in most males, but one excep-
tional specimen has a ratio of 0.13!
Remarks and Range. T.c.melanura appears to be an extremely local
race found only in the vicinity of Mafil in central Valdivia. It has
undoubtedly been derived from c. chilensis, which it resembles very
closely, by a greater production of melanin. But why there should be
such a local melanistic race is hard to say, for the entire region of
southern Chile has a marine climate and typical chilensis is not only
found along the coast of Valdivia but also as far south as Puerto Montt
in Chiloe.
This race has progressed a bit farther than c. chilensis in the evolu-
tion of the color pattern from spotted to lineate, for there is no recapit-
ulation of the spotted type. The late embryo removed from C.M.N.H.
3878 is light brown with a lighter middorsal line bordered by dark
lines, and a dark lateral line on each side. These dark lines are com-
plete and not formed by small fusing spots as in young c. chilensis.
DISCUSSION
Relationship of the Described Forms
Although the large series of Tachymenis discussed above readily
separates into three species complexes, there can be no doubt but that
they form a compact phylogenetic group. T. peruvian a seems to be
the most primitive type and therefore the one closest to the ancestral
stock. Such an ancestral stock probably occupied the Andes of central
and southern Peru where penwiana is now most abundant. The source
of this stock cannot even be guessed at since the relationships of the
various genera of South American Boiginae is so poorly known. Per-
haps it evolved from a lowland form which worked its way up into the
mountains, or perhaps from a form which followed the Andean range
south from some northern point. In any case, given an ancestral stock
resembling peruviana and occupying the Andes of centml and southern
Peru, lines of evolution to the other forms can be traced. These, how-
ever, are necessarily hypothetical.
Probably ajJinis was the first offshoot. It evolved in Peru primarily
by a reduction of the number of scale rows from 19- 19-15 to 17-17-15.
This was accompanied by a marked reduction of the conspicuous
peruvia.?/a color pattern, but not a complete loss (cf. figs. 7 and 12).
Judging by the present rarity of ajJinis, it was a small branch, and the
remaining portion of the stock stayed more or less unchanged as
peru viana. The separation of ajJiuis was complete. No ajJil1is-pcruviana
intergrades are known e\'en though both forms occupy the Andean
region of central and southern Peru. They have even been taken in the
same valley (Urubamba) although not at the same locality.
T. peruviana spread widely to all parts of the Andes of central and
southern Peru, and into those of Bolivia. It even worked down from
the highlands into some of the lower valleys. It has been taken at Vitor
(1,630 meters) on the west side of the Peruvian mountains, and on the
east side in the department of Madre de Dios in Peru and the city of
Santa Cruz (c. 500 meters) in Bolivia. Despite this great spread, it has
remained more or less intact. Local populations differ slightly of
course, but the differences are widely scattered and not arranged in
any clinal sequence. Apparently peruvian{t can reach high enough
elevations to cross mountain passes and allow the various populations
to intermingle to a certain extent. The capture of a number of speci-
mens on the upland plateaus at nearly 4,000 meters elevation where the
vegetation is distinctly alpine indicates that this is true. Most speci-
mens, however, prefer the region of grassy steppes at elevations which
are a bit lower.
But despite this ability to intermingle, peruviana did give rise to a
small offshoot - tarmellsis - at the extreme northem part of its
range. T. tarmensis is known only from the vicinity of Tarma in J unin.
Its scparation from peruviana is not as well marked as in ajJinis. [ts
evolution has been a function of an increase in the number of solid
maxillary teeth from a maximum of 10 in pcruviana to 12, loss of the
apical pit in the dorsal scales which is already poorly developed in cer-
tain peruvian a populations, a slight increase in the number of caudals
(female peruviana maximum is 50; female tarmel1sis 55), and a complete
loss of the marked peruvialla dorsal color pattern so that the snake is a
uniform light greyish brown above (d. figs. 5 & 7, 10 & 11). It is
possible that tarmeusis is a race of peruvian a, but this can not be de-
cided until more material becomes available.
Despite the differences mentioned above, ajJinis, permiana, and
tarme1lsis have certain features in common as mentioned in the intro-
duction so they have been placed into one complex. Two entirely
separate side lines have evolved from peruvialla in the center of this
complex - attl'lIuata at the edge of the eastern tropical lowlands, and
ehile1lsis along the coast of Chile. .
1'. attenuata appears to have arisen from certain populations of
pel'uviana which migrated to the edge of the tropical lowlands and
became somewhat adapted to the changing conditions by elongating
slightly. As mentioned in the introduction, the elongation of the body
caused an increase in the number of caudals and total ventrals, and in
the ratio of tail length to total length. The separation was made more
complete by a change in the penial characters, and an increase in the
number of solid maxillary teeth. This has been pointed out above.
The color pattern of the ancestral peruviana remained to a great extent.
T. attenuata is now found along tlle edge of the tropical lowlands of
southern Peru and northern Bolivia. There was probably only one
uniform species at first, but du ring the course of time the Peruvian
and Bolivian populations have diverged a bit in their color pattems.
T.a.attcnuata in Peru masked the typical dorsal pattern by the develop-
ment of a profuse black flaking on the back, while in a. boliviana the
rows of dorsal black spots fused at their corners to give a checkered
effect (d. figs. 5-7 & 13-16). Minor differences in the number of teeth
and number of abdominals and caudals also appeared.
One paradox presents itself in connection with this theory of the
evolution of attenuata. That is, if attenuata arose from certain peruvian a
populations when they migrated down to the edge of the eastern tropi-
cal lowlands, how is it that other populations have reached quite low
elevations in this region without changing significantly? For example,
both forms have been taken in the Peruvian department of Madre de
Dios. In Bolivia peruviana has been taken at the city of Santa Cruz
(c. 500 meters), and attenuata in the department of Cochabamba at an
elevation of 2,500 meters! The problem Can not be definitely answered
without a better idea of the environment in which the specimens were
taken, but it can be pointed out that there are mountains in parts of
Madre de Dios, that the tropical lowlands reach a high elevation in
Cochabamba
1
, and that Santa Cruz, despite its low elevation, is adja-
cent not only to the Amazonian forests but also to a grazing and agri-
cultural region. Thus the abo\'e concept of the evolution of attenuata
along the edge of the tropical lowlands of the Amazon Basin is not
necessarily thrown out despite a conflict in elevations.
T. chilensis, the second marked sideline arising from peruviana
stock, evolved when that form spread down the western side of the
Andes to thc coast of Chile. This sideline also became distinct from
peruviana in such features as penial characters, oculars, labials, and
coloration as pointed out in the introduction. As mentioned, the penis
resembles that of attenuata, but this must represent a case of parallel
evolution since the two are completely isolated from one another by
the Andes. The southward migration of Tachyrnellis into Chile must
havc taken place some time ago when 'conditions in northern Chile
were not so extremely arid. Subsequently a hiatus appears to have
developed in this region. At least no specimens have been obtained
there, but more collecting is necessary to be certain.
As chilens;s spread southward along the coast, three races appeared
more or less in north-south sequence. These differ from one another
in the number of abdominals and caudals, and in coloration. The num-
ber of abdominals and caudals increases from north to central Chile,
and then falls off a bit in certain parts of the south. This cline shows
up best in the number of total ventrals as diagrammed on page 4l.
The color difference is even more marked, and striking evolutionary
tendencies can be seen (figs. 20- 22). T. c.assirniiis, the most northern
race, typically has a pattern which approaches that of peru viana.
Both forms have a narrow light middorsal line or row of small spots
which is bordered by a row of very obvious large black spots. Laterally,
on the fourth scale row, each has a series of smaller dark spots which
I I\laps of the American Geographic Society show "L1i.S Yungas" reaching an elevation of at
least 3,000 meters in Cochabamba.
is tending to fuse into a line (cf. figs. 6 & 7 with 20). T.c.a8similia
differs in that the rows of dorsal spots fuse into lines posteriorly.
Thus assimilis has the spotted appearance of peruviana anteriorly,
while posteriorly the pattern is more !ineate.
220
210
T
200
190
...L
180
c. assim ilis
T
c. chilensis
r
I
c. melanura
In c. chilensis, this tendency is carried a step further. The light
middorsal line is very pronounced, the large bordering black spots are
reduced to very small ones which fuse into a very narrow line for the
entire length of the animal, and the lateral line is accentuated by a
lightening of the ground color above it (fig. 21). Thus c. chilensis has a
conspicuous lineate pattern quite distinct from the spotted one of
peru"iana but easily derived from it. In fact there is a bit of ontogene-
tic recapitulation, for the young of c. chiZellsis tend to be more spotted
than the adul t8.
T.c.me/allma is the most recent race. It has evolved from c. chiZensis
by developing a great deal of dark pigmentation. This tends to ob-
scure the typical c.chiirnsis pattern, but traces of it can be seen in the
adults (fig. 22). The young, judging from a late embryo, bave lost all
trace of a spotted pattern and merely recapitulate the adult pattern of
c. chilcllsis. At present mcianura is a local race known only from tbe
vicinity of }Iafil in Yaldivia. Typical chilensis is found to the north
and south of it. The climate of this region is marine, and the recently
evolved melanura would appear to be better suited for such condi-
tions. If so, meZallura would be expected to gradually replace chilellsis
in the forested south.
By way of summarizing the "bo\e relationships, a tentative phylo-
genetic tree is presented below:
~
complex
c. melanura
c. chileLi8 chilensis complex
peruviana peruviana complex
Offi\j
Ancestral stock
resembling peruviana
T[(chymenis migrated from Peru and Bolivia down the west side of
the Andes to the coast of Chile, and here gave rise to chilensis. It is
absent in the higher mountains of Chile, but this is not surprising when
one considers the ruggedness and severity of climatic conditions in the
higher parts of the Chilean Andes. But one problem remains. Why
has there not been a similar migration down the east side of the moun-
tains into Argentina? The scrub woodlands and grass steppes of the
region should be ideal for Tachymenis, yet it is conspicuous by its
absence.! There is the possibility of course that more collecting would
reveal its presence. A real absence would be hard to explain unless
in terms of lack of time, or occupation of its ecological niche by other
forms. l'.fore field work is necessary.
Outlying Species
All the snakes discussed above form a natural phylogenetic unit.
But in addition to these forms centering around pcruviana, three other
species have been added to the genus since Boulenger and others limited
it to include only pcruviana and its allies. These are elongata Despax
(1910) from Tablazo de Paita, Peru; brasiliensis Gomes (1918) from
Pindamonhangaba in the State of Sao Paulo, Brazil; and surinamcnsis
Dunn (1922) from Surinam. Their relationship to peruviana and allies
is a problem because of certain morphological differences and their
outlying geographic position. Perhaps they spread peripherally from
the Andes at some distant time, and a large hiatus has since developed;
perhaps they and pcruviana had some common lowland ancestor, or
perhaps they have been misassigned to Tachymenis. Dunn (1922) says
of surinamel1sis, "I am in some doubt as to whether the generic assign-
ment of this snake is correct. . .. It is evidently closely allied to
Tachymenis eZongata." C'.10rphological characters should eventually
throw some light upon this problem, but the status of these snakes must
remain questionable until their anatomy and range is better known.
A reexamination of the types and accumulation of more material is
necessary. Existing descriptions omit certain important features such
as dentition, body proportions, penial characters, etc.
1 Amaral (1930) gives the range of peruviana (of Boulanger) as including Argentina and Para-

proved to be Dryophaylax. Other reports in the literature of the occurrence of perulI'iana in such
places as Ecuador have also result ed from misidentification.
Redefinition of Tachymenis: Synonymy, Morphological
Characters, Range, Habits
Genus TACHYMENIS
1835 Tachyrrumis Wiegmann, N. Acta. Ac. Leop.-Carol., 17, p. 252, pI. 20,
fig. 1 (type peruviana).
1837 Coro';ella Schlegel (not of Laurenti), Phys. Serp., 2, p. 70, (part).
1843 Ophis Fitzinger (not of Wagler), Syst. Rept., pt. I, p. 28, (part).
1854 Dipsas Dumeril and Bibron (not of Laurenti), Erpet. Gen., 7, p. 1159,
(part).
1863 Mesote. Jan, Arch. Zool. Anat. Fisiol., 2, p. 308, (part).
1863 P.amrnophylax Jan (not of Fitzinger), Arcb. Zool. Anat. Fisiol., 2, p.
311, (part).
1915 Leirnadophis Barbour (not of Fitzinger), Proc. BioI. Soc. Washington,
28, p. 149.
Since peruviana is the type of Tachymenis, peruviana and its imme-
diate allies form the nucleus of the genus. It may be that they con-
stitute a subgenus or perhaps the entire genus. Consequently a knowl-
edge of the questionable outlying forms is not absolutely necessary for
an understanding of the characters which must be used to define the
genus. It can be described in terms of the least common denominators
of affinis, peruvial1a, tarmensis, attenuata, and chilensis.
1
All are snakes of moderate size. The largest individuals are only
about 26 inches long. The ratio of tail length to total length is between
0.14-0.18 in the females of most of the forms, and 0.15-0.21 in most
males. Some forms, however, have a relatively longer tail so the ratio
is increased to 0.23 in females and to 0.24 in males. The body propor-
tions of most of the snakes are moderately stocky. A few forms are a
bit more elongate. An oval shaped head is only;<slightly distinct from
the neck. The body of all is more or less cylindrical in section. The
eye is of moderate size with a subeliptical shaped pupil.
Their hemipenis is bifurcate, although not deeply so, with a bifurcate
ductus spermaticus. The tip is spinose becoming calyculate in some.
The body is spinose in all, but the length and distribution of these
spines vary from species to species. The base of all forms bears a few
1 [n discussing the range of variation of such Quantitative characters as number of teeth and
scales, I have only used my observations and those of the original authors in the type descrip-

unreliable.
WALKER: THE SNAKE, TACHHIENIS PERUVIANA WIEGMANN 45
scattered minute spines. Some have several very large basal spines
as well.
The solid maxillary teeth are subequal or increase a bit in length
posteriorly. Their number ranges from 5- 16. These are followed after
a gap by a pair of large, distinctly grooved fangs. The first few
mandibular teeth are quite small, but the length increases rapidly,
reaches a maximum in the front of the jaw, and then decreases grad-
ually.
All the typical cephalic plates are present. In all, the rostral is
broader than deep, slightly visible from above; internasals a bit shorter
than the prefrontals; frontal 1)1' to 2 times as long as broad, longer
than its distance from the end of the snout, as long as or shorter than
the parietals; supraocular of normal size; nasal generally entire, occa-
sionally semidivided; preoculars 1 or 2, rarely 3, the top preocular
reaching the top of the head but not contacting the frontal; postoculars
typically 2, rarely 1 or 3; temporals usually 2- 2 or 2-3 but many vari-
ations are common; supralabials either 7(3-4) or 8(4- 5), departures
from these are rare; infralabials variable but generally 9(4) or 9(5);
anterior pair of chin shields as long as or longer than the posterior.
The dorsal scales are smooth and generally have a single apical pit,
although this is poorly developed in some and completely absent in other
forms. One individual had indications of two pits. The scales are
arranged in an odd number of parallel rows with a reduction pattern of
19-19-15 or 17-17-15. Only a small amount of variation is encoun-
tered in the number of cervical or preanal rows.
The abdominals are rounded and not extremely numerous. The
female range is 134-158; males 136- 169. The anal is divided except
in a few aberrant individuals. The caudals are paired in most,
but some have a few entire ones. The female range is 37- 65; male
40-69. The total ventral range in females is 176- 221; males I84- 221.
The color pattern in these snakes is quite diverse, but some common
features can be seen. In the basic pattern, there is a small dark streak
extending from the snout through the eye to the angle of the mouth.
A similar streak runs from the parietals to the angle of the jaw. The
labials are variably marked with black. Most specimens have such a
mark under each eye. There is a light line or series of spots on the
middle of the back. This is bordered on each side by a series of dark
spots which often fuse into a stripe or line. There is a similar lateral
series of spots or line on the fourth row of scales. Most of the dorsal
scales in the majority of forms have a dark apical dot about the size
of and in the position of the apical pit. There is a large number of
irregular dark markings ventrally which may be arranged in a linear
manner. This pattern is quite fundamental and appears in practically
all the forms of Tachymenis. In a couple of forms it is masked by a
secondary black flaking or melanism but traces of the fundamental
pattern can be seen even in these. The pattern is, however, com-
pletely lost in tarmen<lis. The spotted pattern is more primitive than
the lineate, for it is found in the adults which are believed to be
closest to the ancestral type, and in the young of certain of the lineate
forms. The most advanced types such as chilensis melanura have lost
all trace of it- even the young are lineate.
Sexual dimorphism is manifest in the genus in terms of the number of
abdominals and caudals. Oddly enough, the males have a greater
number of abdominals as well as caudals. Consequently there is only
a slight overlap between the number of male and female total ventrals.
This tendency could be seen to a degree in all the forms studied if both
sexes were available. However, it is most apparent in analyses of
single populations. The specimens of peruviana from Huanta illustrate
this point very well. Their counts are summarized below in tabular
form:
total
abdominal caudal ventral
range average range average range average
6 females 146-150 148 41-47 44 187-196 191
6 ma,les " 148- 156 152 49-52 51 200-206 202
The above tendencies are also reflected in the ratio of tail length to
total length which is a bit higher in the males, and in the slightly
greater male size. Sexual dimorphism could not be detected in other
characters.
The genus seems to center in the Andes of the southern half of
Peru and northern Bolivia. I t reaches elevations there as high as 4,000
WALKER: THE SNAKE, TACI!YMENIS PERUVIANA WIEGMANN 47
meters, but is most abundant in the region of grassy steppes at eleva-
tions which are a bit lower. It has extended from the central Andes to
some of the lower valleys on both sides of the mountains. It has even
reached the edge of the tropical lowlands of the Amazon basin, but
has not spread very widely there. It has also spread down the west
side of the Andes to the coast of Chile. It is found at lower elevations
in Chile presnmably becanse of the unfavorable nature of the higher
mountains. lVIembers of the genus are absent in the se\'ere deserts of
northern Chile, but do occur in the semiarid regions. They are prob-
ably found in the valley bottoms in such places where there would be
some vegetation for cover. The snakes are very abundant in the medi-
terranean portion of central Chile where they con,titute one of the
commonest species. They have extended south from here into the
forested and lake region of Chile but have not been taken below Puerto
Montt. The genus apparently has not migrated down the east side of
the Andes to the plains of Argentina.
Notes on the Habits of the Various Forms
Since field notes have seldom been taken, little direct information is
available on the habits of Tachymcnis. it is p03sible to
make a few deductions on the basis of habitat and morphology. In as
much as the members of the prrllviana and chilrnsis complexes are
fairly stocky snakes, and the bulk of the specimens have COme from
alpine, semiarid, or mediterranean regions, thcy must be terrestrial.
Those from southern Chile were taken in forested country, but even
these have the body form of a terrestrial snake. The Montana and
Ynngas range of members of the aitrll1tata complex, together with the
more elongate body form suggests the possibility that the terrestrial
life is supplemented by a semiarborial habit in these forms.
The subeliptical shaped pupil of all of these snakes may indicate
that they are most active at night in their search for food, etc. Exam-
ination of the stomach contents of ten specimens that had obviously
eaten recently re\'caled that their diet consists primarily of small frogs,
but occasionally toads and terrestrial lizards are captured. Their
rear fanged poison apparatus would help to subdue the more active
prey. The stomach contents of the specimens examined is tabulated
below:
Species Specimen Locality Stomach Contents t
T.p. peruuiana M.C.Z. Yanama, Junin, Peru Remains of a frog
45916
T.p. peruviana C.M.N.H. Huanta, Ayacucho, Bufo spinulosus
39647 Peru trifolium Tschudi
T.p. peruviana C.M.N.H. JuIiaca, Puna, Peru
Liolaemus
34086 multiformis
multiformis
(Cope)
T. attenuata A.M.N.H. Incachaca, Cochabamba,
Remains of a
boliviana
36017 Bolivia
frog and lizard
T.c. chilensis
C.M.N. H. Valparaiso, Aconcagua,
Eupsophus
40117 Chile
taeniatus (Girard)
T.c. chilensis
A.M.N.H. Coraeol, Concepcion,
Eupsophus
18328 Chile
taeniotus (Girard)
T.c. chilensis
C.M.N.H. Curacautio, Cautin,
Pleurodema
6221 Chile
bibroni Tscbudi
'P.c. chilensis C.M.N.H. Traiguen, Cautin, Chile
Liolaemus pictus
31617
pictus (Dum6ril
and Bibron)?
T.c. melanura C.M.N.H. Malil, Valdivia, Chile
Remains of a frog
3867
T.c. melanura C.M.N.H. Malil, Valdivia, Chile
Eupsophu8
3875
taeniatus (Girard)
lI.amindebted to J\Ir. B. Shreve for the identification of certain of the stomach content
specimens.
WALKER: THE S:>IAKE, TACHnIE:>I IS PERUVl.\ XA WIEmlAN:>I -19
No information is m"ailable on the reproduction of aitl!1lUata or
offinis, but examination of the oviducts of adult females' of peruviana
al)(l chilensis show that these forms are ovoviviparous and give birth
to from three to fourteen' young. Fertilization takes place in peruvian a
about May and the young are born in September. The material on
chilo1sis is not as conclusive, but seems to indicate that the reproduc-
tive c,rcle extends from November to February or perhaps even later.
The data is tabulated helow:
Tachyrnenis pCT1Ivialla:
Condition of the
Dale Specimen Locality Emhryo
January .. . ..
February . . . , .
March C.J\1.N.H. Haunta, Ayachucho, Oviducts empty
39650 Peru
M.C.Z. Pachacayo, Junin, Oviducts empty
42435 Peru
April . . . . .
May M.C.Z. Yauyos, Lima, Peru 4 eggs present; em bryo
45914 not. visible grossly
..
1 C.M.N.H. Yunguyo, Puna, 10 eggs present; embryo
40179 Peru not visible grossly
" 1 C.M.N.H. Yunguyo, Puna, 5 eggs present; embryo
" 1 C.M.N.H. Yunguyo, Puna, 5 eggs present; embryo
"
19 C.M.N.H. Tarma, Junin, Peru Oviducts empty
5698
I All adult females with a known eollecting date were examined by palpation for eggs. The
pregnant ones were opened. and one or more of the embryos studied,
'Fourteen eggs were counted by palpation in U.S. N.M. 56442. This specimen is from Chile
but since no further data is available, it has not been included in the accompanying tables.
50 BULLETIN: MUSEUM OF CO}IPARATIVE ZOOLOGY
Tachymenis peruviana:
Date Specimen Locality
Condition of the.
Embryo
June . .. . ,
July .... .
August 29 C.M.N.H. Pisacoma, Puna, Peru 6 eggs present; immatur
40212 embryo; coiled length
15 mm.
September 14 C.M.N.H. Collacachi, Puna, 6 eggs present; immature
34156 Peru embryo; coiled length
Smm.
"
14 C.M.N.H. Collacachi, PUIlD, 8 eggs present; immature
34160 Peru embryo; coiled length
12mm.
"
16 C.M.N.H. U reo, eusen, Peru Oviducts empty
34105
"
16 C.M.N.H. Urea, Cusco, Pem 3 eggs present; immature
10 mm.
" 19 C.M.N.H. Jnliaca, PunD, Peru Oviducts empty
34086
" 19 C.M.N.H. Urea, CUReo, Perll 3 eggs preseIit; mature
32 mm., uncoiled lOS
mm.
"
19 C. M.N.H. UreD, CUBeD, Peru 8 eggs present; immature
34132
embryo; coiled length
17mm.
October .. . ..
November .. . ..
December . . ...
WALKER : THE SNAKE, TACHYMENI S PERUVI ANA WI EGMANN 51
Tachymenis chilensis:
Condition of the
Date Specimen Locality Embryo
January .....
February 14 C.M.N.H. Mali!, Valdivia, 7 eggs present; embryo
to 28 3868 Chile not visible grossly
C.M.N.H. Mali!, Valdivia, Oviducts empty
3872 Chile
C.M.N.H. Mafil, Valdivia, Oviducts empty
3876 Chile
C.M.N.H. Mafil, Valdivia, 4 eggs present; mature
3878 Chile embryo; coiled length
21 mm., uncoiled 98
mm.
3879 Chile
3883 Chile
March 26 C.M.N.H. Valparaiso, Acon- Oviducts empty
40117 cagua, Chile
March 26 C.M.N.H. Concepcion, Oviducts empty
to April 20 3863 Concepcion, Chile
May C.M.N.H. Chiquahue, Bio Bio, Oviducts empty
23808 Chile
June
.. ".
July .... .
August 9 C.M.N.H. Domeyko, Atacama,
to 16 5771 Chile
September . . .. .
October .... .
November C.M.N.H. Traignen, Cautin, 8 eggs present; embryo
31617 Chile not visible grossly
December ... ..
52 BULLETIN: MUREUM OF COMPARATIVE ZOOLOGY
I t is noteworthy that peruvian a, although occurring a bit south of the
equator, has the reproductive cycle characteristic of snakes of the
northern hemisphere. This is not too surprising when it is realized
that its remote ancestors probably came from the north, and that the
seasonal differences near the equator are not pronounced enough to
require a change. However, the breeding period of chilensis, or at least
the southern members of the species, has shifted to conform to the
summer season in southern South America. It would be of interest to
know whether this change has become a specific difference, or whether
it is limited to the southern populations of chilensis. This cannot be
decided until more material becomes available from northern Chile.
Key to PERUVIANA and Allies
As pointed out above, Tachymcnis peruviana and its immediate
allies form at least the nucleus of the genus. A key to their identifica-
tion follows:
Body form moderately stocky. Correlated with this, the caudals
are under 60 and thc ratio of tail length to total length is under 0:22.
Solid maxillary teeth 6- 12 .................................. 2
11 Body form elongate. Correlated with this, the caudals are 60 or
over and the ratio of tail length to total length is 0.22 or over.
Solid maxillary teeth 12-16. Found in southeastern Peru or eastern
BoI'ivia along the edge of the tropical lowlands ............... .
The attenuata complex ...... 5
2 (1) Supralabials 8(4-5); preocular 1; body of hemipenis covered
with spines of subequal size, no large basal spines. Found in the
Andean region of Peru and Bolivia .......................... .
The peruviana complex ...... 3
21 Supralabials 7(3-4); preoculars 2; body of the hemipenis covered
with spines which increase in size posteriorly, several large basal
spines. Found along the coast of Chile . .. .. . . . ... . ........ ... .
The chilcnsis complex ...... 6
3 (2) Midbody scale rows 19 ...... . . .. .. . . . . .. ............. . . 4
3
1
Midbody scale rows 17. Found in the Andean region of Central
and Southern Peru ..... . ............ . .... . ............ affinis
4 (3) Dorsal scales with single apical pits although these are some-
times poorly deyeloped, a well developed spotted dorsal ground
color, female caudals 50 or under, 6- 10 solid maxillary teeth.
Found in the Andean region of central and southern Peru ..... . .
pcruviana
4
1
Dorsal scales without apical pits, Jorsal color a uniform grayish
brown with no trace or only a very slight trace of darker spots,
female caudals oYer 50, 12 solid maxillary teeth. Found in or near
Tarma, Department of Junin, central Peru ........... . tarmensis
5 (11) Dorsal scales generally heavily flaked with black, obscuring
any other pattern; 12-14 solid maxillary teeth. Found in south-
eastern Peru .......... . ............. . ...... attenuata attenuata
51 A checkered dorsal color pattern, 14-16 solid maxillary teeth.
Found in eastern Bolivia ...... . ..... . ....... attenuata boliviana
6 (21) A distinctly spotted or lineate dorsal color pattern .......... 7
6
1
Dorsal ground color so melanistic that an underlying lineate pattern
is nearly completely obscured. Found in or near l\fafil, Province of
Valdivia, southern Chile ..................... chilensis melanura
7 (6) The rows of dorsal spots on either side of the light middorsal line
are very prominent being two or more scales wide. The spots fuse
into a stripe posteriorly, bnt are usually distinct anteriorly. Found
in northern Chile from the province of Atacma to the province of
Santiago ................... ... .. . . .. ....... chilensis assimilis
7
1
The rows of dark dorsal spots are much reduced in size and fuse to
form a line. They are not over one scale wide. Found in central
and southern Chile from the province of Aconcagua to the province
of Chiloe ......... , .......... . .............. chilensis chilensis
Relationship between Tachy,nenis and other Boiginae
I conclude this discussion of Tachymenis by emphasizing how little
we know about its relation to other genera of Boiginae in South Amer-
ica. At one time there was confusion between Tachymenis and Ery-
throlamprus. Boulenger and others decided that many spccies origi-
nally described as Tachymenis were misassigned and that they should
be referred to Erythrolamprus. A list of these can be found in the intro-
duction. Boulenger pointed out in his Catalogue (1896) that members
of Erytholamprus differ in having fangs which are only feebly enlarged
and sometimes not grooved, a round pupil, no scale pits, a variable
number of scale rows (15, 17, 19, 21, or 23), and ventrals which are
sometimes obtusely angulate laterally. These criteria should separate
the two genera, but the situation has become confused because it has
been realized that Erythrolamprus as redefined by Boulenger is an
omnium gatherum. Among other things, it included all of COlliophanes
(Cope, 1860) which has since been removed from the synonymy.
Dunn (1922) feels that Coniophanes is very close to Tachymenis, and
perhaps inseparable. Tachymenis is the older name.
BIBLIOGRAPHY
AMARAL, A. DO
1925. "South American Snakes in the Collection of the United States
National Museum." Proc. U. S. Nat. Mus., 67, Art. 24, pp. 1-30.
1930. "Estudos sobre Ophidios Ncotropicos. XVIII - Lista remissiva
dos Ophidios da Regiao Neotropico." Mem. lnst. Butantan Sao
Paulo, 4(1929), pp. 129-271.
AMERICAN GEOGRAPHICAL SOCIETY OF NEW YORK.
1922-1938. Peruvian Sheets of the "Map of Hispanic America,"
1:1,000,000. .
1942. South American Sheets of the "Map of the Americas," 1 :5,000,000.
BARBOUR, T.
1915. "A new Snake from southern Peru." Proc. BioI. Soc. Washington,
28, pp. 149-150.
BARBOUR, T., and NOBLE, G. K.
1921. "Amphibians and Reptiles from southern Peru collected by the
Peruvian Expedition of 1914-15 under the Auspices of Yale Uni-
versity and the National Geographic Society." Proc. U. S. Nat.
Mus., 58, pp. 609--{l20.
B1NGHoAM, H.
1916. "Further Explorations in the Land of the Incas. The Peruvian
Expedition of 1915 of the National Geographic Society and Yale
University." Nat. Geog. Mag., 29, pp. 413-473, many figs.
BOULENGER, G. A.
1893-1896. "Catalogue of the Snakes in the British Museum of
Natural History," 1(1893), 2(1894), 3(1896). London.
BOWMAN, I.
1916. "The Andes of southern Peru." Amer. Geog. Soc. New York.
CARLSON, F. A.
1943. "Geography of Latin America." New York.
DES PAX, M. R.
1910. "Mission Geodesique de l'Equateur. Collections recueillies par
M. Ie Dr. Rivet. Listc des Ophidicns et Description des Especes
nouvelles." Bull. Mus. Hist. Nat. Paris, 16, pp. 368-376.
DUNN, E. R.
1922. "Two new South American Snakes." Proc. BioI. Soc. Washington,
36, pp. 219-20.
GOMES, J. F.
1918. "Contribuiyao para 0 Conhecimento dos Ophidios do Brazil.
Pt. II. Descri,ao de duas Especies novas." Mem. lnst. Butantan
Sao Paulo, 1(1), pp. 78-83, pI. 14, figs. 1-2.
WALKER : ;rHE SNAKE, TACHYMENIS PERUVIANA WIEGMANN 55
GRIFFIN, L. E.
'1916. "A Catalogue of the Ophidia from South America at present con-
tained in the Carnegie 1Iuseum, with Descriptions of some new
Species." Mem. Carnegie Mus., 7, pp. 163-228, pl. 28.
HELLMICH, 'V.
1938. "Anotaciones para el Conocimiento de las Culebras de Chile."
Rev. Chilena Hist. Nat. Santiago, 41, pp. 107-110.
JAN, G.
1863. "Enumerazione Sistematica degli Ofidi appartenente aI Gruppo
Coronellidi." Arch. Zool. Anat. Fisiol., 2, pp. 213'::'330.
1866. "Iconographie general des Ophidiens", 18, 6 pIs.
MEYEN, F. J. F.
1834. "Reise urn die Erde," 1. Berlin.
RUDOLPH, W. E.
1929. "The new Territorial Divisions of Chile with Special Reference to
Chil6e." Geog. Rev., 19, pp. 61-77, 16 figs.
SCHLEGEL, H.
1837. "Essai sur la Physionomie des Serpens," 2. Amsterdam.
SCHMIDT, K. P., and WALKER, W. F., JR.
1943. "Peruvian Snakes from the University of Arequipa." Zool. Ser.
Field Mus. Nat. Hist., 24, pp. 279-296.
STIGLICH, G.
1922-1923. "Diccionario Geographico del Peru." Lima.
\VEBERBAUER, A.
1936. "Phytogeography of the Peruvian Andes," in MacBride, J. F.,
"Flora of Peru." Bot. Ser. Field Mus. Nat. Hist., 13, pt. 1, pp.
13-37, map.
WERNER, F.
1898. "Die Reptilien und Batrachier der Sammlung Plate." Zool.
Jahrb. Sup., 4, pp. 244-78, 2 text figs., pIs. 13-14.
1901. "Reptilien und Batrachier aus Peru und Bolivien." Abhandl.
und Berich. Kon. Zool. Anthrop.-Ethnog. Mus. Dresden, 9(2), pp.
1-14,8 text figs.
1904. "Ergebnise Hamburger Magalh. Sammelreise." N aturhistorishes
Mus. zu Hamburg, 1, Rept. und Batr.
W IEOMANN, A. F. A.
1835. "Beitrage zur Zoologie, Gesammelt auf einer Reise urn die Erde
von F. J. F. Meyen." Ahhandl. 7, Amphibien. N. Acta. Ac. Leop.-
Carol., 17, pp. 183-268, pIs. 13-22.
PLATES
\V ALKER-Tachymenis peruyiana Wiegmann
PLATE 1
Fig. I. Map of southwestern South America showing the approximate
range of the species of Tachymenis studied above. The outline for this map
was obtained through the courtesy of the American Geographical Society. The
key to the map follows. The range of T. tarmensi. is in cent.ral J unin only.
Its symbol may not be clear.
o Tachyrnenis peruIJian..a vViegmann
~ Tachymenis tarmensis Walker.
Tachymenis affini. Boulenger
~ Tachymenis attenuata attenuata Walker
I!!IIl!II Tachymenis attenuata boliviana Walker
_ Tachymenis chilensis assimilis (Jan)
~ Tachymenis chilensis chilensis (Schlegel )
III Tachyme-nis chilensis melanura " "'a1ker
BULL. MUS. CaMP, ZOOL.
W ALKER. TACHYMENIS PERUVIAN" W I EGMANN. PLATE 1
\
. \,
\
~ ~ .
-'.- ___ ----t-----j
I \ I
\
- - ~
PLATE 2
WALKER-Tachymenis peruviana Wiegmann
PLATE 2
Fig. 2. Tachymenis peruviana Wiegmann. Dissection of the hemipenis of a
specimen from Vitor, Arequipa, Peru (C.M.N.H. 34260). x5.
Fig. 3. Tachymeni. attenuata attenuata Walker. Dissection of the hemi-
penis of the type from the department of Madre de Dios, Peru (C.M.N.H.
40071). x4.
Fig. 4. Tachy",eni . chilensis chilensis (Schlegel). Dissection of the hemi-
penis of a specimen from Concepcion, Concepcion, Chile (M.C.Z. 16374). x3.
BULL. MUS. CaMP. ZOOL. W ALKER. T ACHYME:NI S PERUVIANA W IEGMANN. P LATE 2 .
2
3
4
PLATE 3
W ALXER-Tachymeni s peruviana Wiegmann
PLATE 3
Fig. 5. Tachymenis peruviana Wiegmann. Lateral view of the head and
neck of a specimen from Huacullani, Puno, Peru (C.M.N.H. 40213) showing
typical coloration. xl v,.
Fig. 6. Same specimen as in figure 5. Dorsal view of the head and neck.
xlV,.
Fig. 7. Tachymenis peruviana Wiegmann. Typical dorsal coloration as seen
in a specimen from Yunguyo, Puno, Peru (C.M.N.H. 40181). x;J4.
Fig. 8. Tachymenis perumar.a Wiegmann. Dorsal pattern of a specimen
from Santa Cruz de la Sierra, Santa Cruz, Bolivia (A.M.N.H. 8793) showing
fusion of the light middorsal spots into a distinct line. This may be the pattern
of dorsalis Werner, 1901. Natural size.
Fig. 9. Tachyrnenis peruviana ,"Viegmann. Dorsal pattern of a specimen
from the department of Madre de Dios, Peru (C.M.N.H. 40075). The row of
light middorsal spots is still in evidence, even though the specimen is very
melanistic. x;J4.
BULL. MUS. CaMP. ZOOL. W ALKER. T ACHYMENIS PERUVIJ\NA W IEGMANN. P LATE 3.
6
7
8
9
PLATE 4
W AL:K:En-TachymeniB peruviana Wiegmann
PLATE 4
Fig. 10. Tachymenis larmensis Walker. Lateral view of the head of the
type specimen from Tarma, Junin, Peru (C.M.N.H. 5698). Notice the loss of
the typical peru,iana pattern. xl7!:!.
Fig. 11. Same specimen as in figure 10. A strip of skin from the back show-
ing the absence of any pattern. x;li.
Fig. 12. Tachymenis ajJini. Boulenger. A strip of skin from the middle of
the back of a specimen from Macchu Picchu, Cuzco, Peru (U.S.N.M. 60721)
showing reduction of color pattern. x%:.
Fig. 13. Tachymenis attenuata attenuata Walker. Lateral view of the head
and neck of the type from the department of Madre de Dios, Peru (C.M.N.H.
40071). xl7!:!. .
Fig. 14. Same specimen as in figure 13. Dorsal view of the head and neck.
xl7!:!.
Fig. 15. Same specimen as in figure 13. Dorsal pattern. x%:.
Fig. 16. Tachymenis attenuata boliviana Walker. Dorsal pattern of the type
from Incachaca, Cochabamba, Bolivia (A.M.N.H. 36020). Natural size.
BULL. MUS. COMPo ZOOL. Wl\l KER. T ACHYMENI S PERUVIANA W IEGMANN, P LATE 4 .
10
1/
12
13
14
15
16
PLATE 6
WALxER-Tacbymenis peruviana Wiegmann
PLATE 5
Fig. 17. Tachymenis chilensis chilensis (Schlegel). Typical dorsal pattern as
seen in a specimen from Talcaguano, Concepcion, Chile (M.C.Z. 2540). x%:.
Fig. 18. Tachymenis chilensis chilensis (Schlegel). Dorsal pattern of another
specimen from Talcaguano, Concepcion, Chile (M.C.Z. 2066) showing a great
accentuation of the typical lineate pattern. x%:.
Fig. 19. Tachymeni. chilensi. chilensi. (Schlegel). Dorsal pattern in a speci-
men from Concepcion, Concepcion, Chile (M.C.Z. 16374) showing a reduction
of the typicallineate pattern. x%:.
Fig. 20. Tachymenis chilensis assimilis (Jan). Dorsal view of the anterior
part of a specimen from Valparaiso, Aconcagua, Chile (A.l\1.N.H. 36147)
showing the large series of dorsal spots beginning to merge into stripes. x%:.
Fig. 21. Tachymenis chilensis chilensis (Schlegel). Dorsal view of the
anterior part of a specimen from Talcaguano, Concepcion, Chile (M.C.Z. 2540)
showing typical pattern of small dorsal black spots merging into a line. x%:.
Fig. 22. Tachymenis chilensis melanura Walker. Dorsal view of the anterior
part of the body of a paratype from Mati!, Valdivia, Chile (C.M.N.H. 3883)
showing a slight retention of the typical chilensis pattern, despite the melanistic
nature of the race. x%:.
BULL. MUS. CaMP. ZObL. W ALKER. TACHYMENIS PERUVI ANA WI EGMANN. PLATE 5.
17
18
19
21
20
22

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